Blend Of Herbivore-induced Plant Volatiles Research Articles

Identity Matters: Multiple Herbivory Induces Less Attractive or Repellent Coffee Plant Volatile Emission to Different Natural Enemies.

Co-infestations by herbivores, a common situation found in natural settings, can distinctly affect induced plant defenses compared to single infestations. Related tritrophic interactions might be affected through the emission of changed blends of herbivore-induced plant volatiles (HIPVs). In a previous study, we observed that the infestation by red spider mite (Oligonychus ilicis) on coffee plants facilitated the infestation by white mealybug (Planococcus minor), whereas the reverse sequence of infestation did not occur. Here, we examined the involvement of the jasmonate and salicylate pathways in the plant-mediated asymmetrical facilitation between red spider mites and white mealybugs as well as the effect of multiple herbivory on attractiveness to the predatory mite Euseius concordis and the ladybug Cryptolaemus montrouzieri. Both mite and mealybug herbivory led to the accumulation of JA-Ile, JA, and cis-OPDA in plants, although the catabolic reactions of JA-Ile were specifically regulated by each herbivore. Infestation by mites or mealybugs induced the release of novel volatiles by coffee plants, which selectively attracted their respective predators. Even though the co-infestation by mites and mealybugs resulted in a stronger accumulation of JA-Ile, JA and SA than the single infestation treatments, the volatile emission was similar to that of mite-infested or mealybug-infested plants. However, multiple infestation had a negative impact on the attractiveness of HIPVs to the predators, making them less attractive to the predatory mite and a repellent to the ladybug. We discuss the potential underlying mechanisms of the susceptibility induced by mites, and the effect of multiple infestation on each predator.

Plant communication across different environmental contexts suggests a role for stomata in volatile perception.

Plants can detect herbivore-induced plant volatiles (HIPVs) from their damaged neighbours and respond by enhancing or priming their defenses against future herbivore attack. Plant communication and defense priming by volatile cues has been well documented, however, the extent to which plants are able to perceive and respond to these cues across different environmental contexts remains poorly understood. We investigated how abiotic changes that modulate stomatal conductance and/or defense signalling affect the ability of maize plants to perceive HIPVs and respond by priming their defenses. During light exposure, when stomata were open and conditions allowed for defense signal biosynthesis, the individual compounds indole and (Z)-3-hexenyl acetate primed maize defenses. Neither compound primed defenses under environmental conditions that closed stomata and/or altered defense signalling. Moreover, plants were not primed when exposed to indole or (Z)-3-hexenyl acetate in darkness (while stomata were closed) and then subjected to simulated herbivory in the light, to ensure defense induction. The full blend of HIPVs primed maize defenses in light conditions but suppressed defense induction during dark exposure and wounding. These findings indicate that environmental context is important for plant communication and defense priming and suggest that stomata play a role in plant perception of HIPVs.

Priming Seeds with Indole and (Z)-3-Hexenyl Acetate Enhances Resistance Against Herbivores and Stimulates Growth.

A striking feature of plant ecology is the ability of plants to detect and respond to environmental cues such as herbivore-induced plant volatiles (HIPVs) by priming or directly activating defenses against future herbivores. However, whether seeds also respond to compounds that are common constituents of HIPV blends and initiate future plant resistance is unknown. Considering that seeds depend on other environmental cues to determine basic survival traits such as germination timing, we predicted that seeds exposed to synthetic constituents of HIPV blends would generate well-defended plants. We investigated the effect of seed exposure to common volatiles on growth, reproduction, and resistance characteristics in the model plants Arabidopsis thaliana and Medicago truncatula using herbivores from two feeding guilds. After seed scarification and vernalization, we treated seeds with one of seven different plant-derived volatile compounds for 24h. Seeds were then germinated and the resulting plants were assayed for growth, herbivore resistance, and expression of inducible defense genes. Of all the synthetic volatiles tested, indole specifically reduced both beet armyworm growth on A. thaliana and pea aphid fecundity on M. truncatula. The induction of defense genes was not affected by seed exposure to indole in either plant species, indicating that activation of direct resistance rather than inducible resistance is the mechanism by which seed priming operates. Moreover, neither plant species showed any negative effect of seed exposure to any synthetic volatile on vegetative and reproductive growth. Rather, M. truncatula plants derived from seeds exposed to (Z)-3-hexanol and (Z)-3-hexenyl acetate grew larger compared to controls. Our results indicate that seeds are sensitive to specific volatiles in ways that enhance resistance profiles with no apparent costs in terms of growth. Seed priming by HIPVs may represent a novel ecological mechanism of plant-to-plant interactions, with broad potential applications in agriculture and seed conservation.

Eliciting Plant Defenses Through Herbivore-Induced Plant Volatiles’ Exposure in Sweet Peppers

Insect herbivory activates plant defense mechanisms and releases a blend of herbivore-induced plant volatiles (HIPVs). These volatile compounds may be involved in plant-plant communication and induce defense response in undamaged plants. In this work, we investigated whether the exposure of sweet pepper plants to HIPVs [(Z)-3-hexenol, (Z)-3-hexenyl acetate, (Z)-3-hexenyl propanoate, (Z)-3-hexenyl butanoate, hexyl butanoate, methyl salicylate and methyl jasmonate] activates the sweet pepper immune defense system. For this, healthy sweet pepper plants were individually exposed to the each of the above mentioned HIPVs over 48 h. The expression of jasmonic acid and salicylic acid related genes was quantified. Here, we show that all the tested volatiles induced plant defenses by upregulating the jasmonic acid and salicylic acid signaling pathway. Additionally, the response of Frankliniella occidentalis, a key sweet pepper pest, and Orius laevigatus, the main natural enemy of F. occidentalis, to HIPV-exposed sweet pepper plants were studied in a Y-tube olfactometer. Only plants exposed to (Z)-3-hexenyl propanoate and methyl salicylate repelled F. occidentalis whereas O. laevigatus showed a strong preference to plants exposed to (Z)-3-hexenol, (Z)-3-hexenyl propanoate, (Z)-3-hexenyl butanoate, methyl salicylate and methyl jasmonate. Our results show that HIPVs act as elicitors to sweet pepper plant defenses by enhancing defensive signaling pathways. We anticipate our results to be a starting point for integrating HIPVs-based approaches in sweet pepper pest management systems which may provide a sustainable strategy to manage insect pests in horticultural plants.

Defense Suppression through Interplant Communication Depends on the Attacking Herbivore Species

In response to herbivory, plants emit volatile compounds that play important roles in plant defense. Herbivore-induced plant volatiles (HIPVs) can deter herbivores, recruit natural enemies, and warn other plants of possible herbivore attack. Following HIPV detection, neighboring plants often respond by enhancing their anti-herbivore defenses, but a recent study found that herbivores can manipulate HIPV-interplant communication for their own benefit and suppress defenses in neighboring plants. Herbivores induce species-specific blends of HIPVs and how these different blends affect the specificity of plant defense responses remains unclear. Here we assessed how HIPVs from zucchini plants (Cucurbita pepo) challenged with different herbivore species affect resistance in neighboring plants. Volatile “emitter” plants were damaged by one of three herbivore species: saltmarsh caterpillars (Estigmene acrea), squash bugs (Anasa tristis), or striped cucumber beetles (Acalymma vittatum), or were left as undamaged controls. Neighboring “receiver” plants were exposed to HIPVs or control volatiles and then challenged by the associated herbivore species. As measures of plant resistance, we quantified herbivore feeding damage and defense-related phytohormones in receivers. We found that the three herbivore species induced different HIPV blends from squash plants. HIPVs induced by saltmarsh caterpillars suppressed defenses in receivers, leading to greater herbivory and lower defense induction compared to controls. In contrast, HIPVs induced by cucumber beetles and squash bugs did not affect plant resistance to subsequent herbivory in receivers. Our study shows that herbivore species identity affects volatile-mediated interplant communication in zucchini, revealing a new example of herbivore defense suppression through volatile cues.

Do Conspecific Herbivores Track Resource Depletion through Host Phenology-Specific HIPVs?

Conspecific herbivores use herbivore-induced plant volatiles (HIPVs) as cues while selecting an optimal site for oviposition. This is to ascertain the availability of nourishment for their progeny so that they get the best chance at survival. In the present study, phytophagous eulophid seed-borer Anselmella kerrichi was significantly (time spent: F 3,40 = 13.12, P < 0.0001; number of entries: F 2,30 = 4.21; P = 0.02) attracted to HIPVs from immature fruits of Java plum, Syzygium cuminii (time spent: 4.77 ± 0.40 min; number of entries: 2.27 ± 0.24) when also given the choice of mature fruits (time spent: 1.76 ± 0.32 min; number of entries: 1.46 ± 0.16), indicating that herbivores can assess re-source depletion from host phenology-specific HIPVs. The chemical cues like α -pinene, β -pinene, (z)-ocimene, undecane, 3,7-dimethyl decane, neo-allo-ocimene, allo-ocimene, ethyl benzoate, 2,6,11-trime¬thyldodecane, α -copaene and β -caryophyllene, which are present in immature fruit volatiles elicited antennal response in a GC-EAD analysis. Olfactometer analyses with the synthetic compounds also revealed that A. kerrichi was significantly attracted to these cues and the synthetic blend composed of the above compounds proved to be much more efficient in attracting female wasps when compared to a natural blend. Field evaluations using the synthetic blend showed that it could attract a significant number of A. kerrichi , indicating the scope of using this blend of synthetic HIPVs as a sustainable IPM tool

Plant volatiles as regulators of plant defense and herbivore immunity: molecular mechanisms and unanswered questions.

Plants release distinct blends of herbivore-induced plant volatiles (HIPVs) upon herbivore attack. HIPVs have long been known to influence the behavior of herbivores and natural enemies. In addition, HIPVs can act as physiological regulators that induce or prime plant defenses. Recent work indicates that the regulatory capacity of HIPVs may extend to herbivore immunity: herbivores that are exposed to HIPVs can become more resistant or susceptible to parasitoids and pathogens. While the mechanisms of HIPV-mediated plant defense regulation are being unraveled, the mechanisms underlying the regulation of herbivore immunity are unclear. Evidence so far suggests a high degree of context dependency. Here, we review the mechanisms by which HIPVs regulate plant defense and herbivore immunity. We address major gaps of knowledge and discuss directions for future mechanistic research to facilitate efforts to use the regulatory capacity of HIPVs for the biological control of insect pests.

Behavioral response of the generalist predator Orius insidiosus to single and multiple herbivory by two cell content-feeding herbivores on rose plants

Multiple herbivory by arthropods with distinct feeding modes often reduces the attractiveness of herbivore-induced plant volatiles to the third trophic level, while herbivory by two species with the same feeding mode yields variable effects. So far, only few studies have examined multiple herbivory with two cell-content feeders. Here, we addressed the effect of multiple herbivory in rose plants by two cell-content feeders, Tetranychus urticae Koch (Acari: Tetranychidae) and Frankliniella insularis (Franklin) (Thysanoptera: Thripidae), on the olfactory preference of the minute pirate bug Orius insidiosus (Say) (Hemiptera: Anthocoridae), a generalist predator, to herbivore-induced plant volatiles (HIPVs). Additionally, we investigated whether the predator’s olfactory preference for HIPVs emitted by rose plants is associated with its feeding preference and prey quality. Y-tube olfactometer assays revealed that O. insidiosus was equally attracted to volatiles emitted by mite- or thrips-infested roses. Although HIPV blends from single-infested and multiple-infested rose plants differed in qualitative and quantitative terms, the minute pirate bug did not discriminate the odors of thrips- or mite-infested plants from multiple-infested plants. Congruent to the olfactory preference toward HIPVs, O. insidiosus did not show preference for either prey species, but consumed larger numbers of spider mites than thrips to complete its development. Therefore, our results showed that multiple herbivory by two cell-content feeders do not change the attractiveness of HIPV emissions compared to those of single-infested rose plants, and that lack of preference of the minute pirate bug to HIPV emissions induced by either spider mites or thrips favors the location of suitable prey.

Birds Bug on Indirect Plant Defenses to Locate Insect Prey.

It has long been thought that most birds do not use volatile cues to perceive their environment. Aside from some scavenging birds, this large group of vertebrates was believed to mostly rely on highly developed vision while foraging and there are relatively few studies exploring bird response to volatile organic compounds. In response to insect herbivory, plants release volatile organic compounds to attract parasitoids and predators of the pests. To test if insectivorous birds use herbivore-induced plant volatiles (HIPV), dispensers emitting a synthetic blend of HIPV typically emitted after insect herbivory were deployed in a maize field along with imitation clay caterpillars. Significantly more imitation insects were attacked by birds when located close to dispensers releasing HIPV than close to dispenser with organic solvent only. Seven times more peck marks, an index of avian predation, were counted on caterpillars in the vicinity of the HIPV dispensers than on insects close to control dispensers. This is the first field demonstration that insectivorous birds cue on HIPV to locate prey in agricultural settings. These results support the growing evidence that foraging birds exploit volatile cues. This more accurate understanding of their behavior will be important when implementing pest management program involving insectivorous birds.

Invasive insect herbivores as disrupters of chemically-mediated tritrophic interactions: effects of herbivore density and parasitoid learning

Invasive insect herbivores have the potential to interfere with native multitrophic interactions by affecting the chemical cues emitted by plants and disrupting the attraction of natural enemies mediated by herbivore-induced plant volatiles (HIPVs). In a previous study, we found that the presence of the exotic herbivore Spodoptera littoralis on Brassica rapa plants infested by the native herbivore Pieris brassicae makes these dually-infested plants unattractive to the main parasitoid of P. brassicae, the braconid wasp Cotesia glomerata. Here we show that this interference by S. littoralis is strongly dependent on the relative densities of the two herbivores. Parasitoids were only deterred by dually-infested plants when there were more S. littoralis larvae than P. brassicae larvae on a plant. Furthermore, the blend of HIPVs emitted by dually-infested plants differed the most from HIPVs emitted by Pieris-infested plants when S. littoralis density exceeded P. brassicae density. We further found that associative learning by the parasitoid affected its preferences: attraction to dually-infested plants increased after parasitoids were presented a P. brassicae caterpillar (rewarding experience) in presence of the odor of a dually-infested plant, but not when presented a S. littoralis caterpillar (non-rewarding experience). A non-rewarding experience prior to the bioassays resulted in a general decrease in parasitoid motivation to respond to plant odors. We conclude that herbivore density and associative learning may play an important role in the foraging behavior of natural enemies in communities, and such effects should not be overlooked when investigating the ecological impact of exotic species on native food webs.

What signals do herbivore-induced plant volatiles provide conspecific herbivores?

Herbivore-induced plant volatiles (HIPVs) have been opined as ‘indirect or direct defenses’ of plants and are extensively studied. In contrast, HIPVs may also indicate that plant defenses have been overcome by herbivores infesting the plant; however, studies on this aspect have so far received little attention. Using the interaction of Capsicum annum (Bell pepper) with its pest Scirtothrips dorsalis (Chilli thrips) as a model system, we studied the role of HIPVs in this selected insect–plant interaction. Multiple-choice olfactometer assays with headspace volatiles collected from different growth stages of un-infested C. annum plants represented by pre-flowering (PF), flowering (FL) and fruiting stages (FR) proved FR volatiles to be highly attractive to S. dorsalis. Further, FR plants were infested with S. dorsalis adults and HIPVs released by infested plants were collected and subjected to multiple-choice olfactometer bioassays. Thrips were significantly attracted to HIPVs than to headspace volatiles of un-infested FR plants or thrips body odour. Coupled GC-EAG with S. dorsalis and HIPVs or FR plant volatile revealed specific compounds that elicited an EAG response. Individual EAG-active compounds were less attractive to thrips, however, synthetic blends of EAG-active compounds at the ratio similar to headspace samples were found to be highly attractive. However, when given a choice between synthetic blends of HIPVs and FR, thrips were significantly attracted to synthetic blend of HIPVs. Our study provides empirical data on signals HIPVs may provide to conspecific herbivores and suggests that the role of HIPVs, mostly generalized as defense, may vary based on the interaction and must be studied closely to understand their ecological functions.

Herbivore-induced plant volatiles accurately predict history of coexistence, diet breadth, and feeding mode of herbivores.

Herbivore-induced plant volatiles (HIPVs) serve as specific cues to higher trophic levels. Novel, exotic herbivores entering native foodwebs may disrupt the infochemical network as a result of changes in HIPV profiles. Here, we analysed HIPV blends of native Brassica rapa plants infested with one of 10 herbivore species with different coexistence histories, diet breadths and feeding modes. Partial least squares (PLS) models were fitted to assess whether HIPV blends emitted by Dutch B.rapa differ between native and exotic herbivores, between specialists and generalists, and between piercing-sucking and chewing herbivores. These models were used to predict the status of two additional herbivores. We found that HIPV blends predicted the evolutionary history, diet breadth and feeding mode of the herbivore with an accuracy of 80% or higher. Based on the HIPVs, the PLS models reliably predicted that Trichoplusia ni and Spodoptera exigua are perceived as exotic, leaf-chewing generalists by Dutch B.rapa plants. These results indicate that there are consistent and predictable differences in HIPV blends depending on global herbivore characteristics, including coexistence history. Consequently, native organisms may be able to rapidly adapt to potentially disruptive effects of exotic herbivores on the infochemical network.

Different population densities and continuous feeding by Oligonychus yothersi (McGregor) (Acari: Tetranychidae) affect the emissions of herbivore-induced plant volatiles on avocado (Persea americana Mill. cv. Hass) shoots under semi-field conditions

ABSTRACTPlants attacked by insects and mites release blends of herbivore-induced plant volatiles (HIPVs), which act as chemical cues that attract predators and/or repel herbivores alongside alerting neighbouring plants. The chemical profiles of avocado (Persea americana Mill. Cultivar Hass) shoots infested with the foliar mite Oligonychus yothersi (Acari: Tetranychidae) were analysed using a gas chromatograph coupled to a mass spectrometer (GC-MS); the different collections of volatiles were performed under semi-field conditions in summer. Four infestation treatments were applied to the avocado shoots (uninfested plant or control; low, medium and high mite-infestation levels). The chemical profiles collected in situ changed qualitatively and quantitatively after infestation, increasing the emissions of (Z)-β-ocimene, (E)-β-ocimene, β-linalool, α-farnesene, farnesyl cyanide and methyl salicylate (MeSA) as opposed to those of the uninfested shoots. The amount of HIPVs emitted by the avocado shoots with the highest O. yothersi population density (250 ± 10 mites per leaf) were greater than those of the shoots infested with the lower population density (60 ± 10 mites per leaf); to the emissions of (E)-β-ocimene, sesquiterpenes and MeSA at higher amounts even after seventeen days of continuous infestation. Likewise, the emissions of HIPVs increased with each infestation level; also fluctuating between collecting dates, which indicates that environmental conditions maximized or constrained those emissions. In addition, the damaged foliar area increased with higher O. yothersi population densities after seventeen days of infestation. These results indicate that higher mite-population densities and continuous O. yothersi-feeding increases the amounts of HIPVs emitted by O. yothersi-infested shoots in summer.

Chemical cues modulating electrophysiological and behavioural responses in the parasitic waspCotesia sesamiae

A better understanding of the chemical cues employed by parasitoids to locate their herbivore hosts will contribute towards effective exploitation of parasitoids as biological control agents against damaging crop pests. The current study was conducted to establish key bioactive compounds mediating behavioural response in the parasitic wasp Cotesia sesamiae (Cameron, 1906) (Hymenoptera: Braconidae) by employing electrophysiological techniques and behavioural studies. Electroantennogram (EAG) recording with synthetic equivalents of herbivore-induced plant volatiles (HIPVs) using C. sesamiae confirmed that 13 compounds elicited electrophysiological activity. In an olfactometer bioassay, only (E)-4,8-dimethyl-1,3,7-nonatriene, (E)-β-farnesene, and (E,E)-4,8,-trimethyl-1,3,7-tridecatetraene elicited behavioural responses from the parasitoid when tested individually at a natural dose, while (R)-linalool and (E)-caryophyllene were attractive at the highest test dose. Other EAG-active HIPVs were not attractive either at lower or higher concentrations, suggesting their role as blend components rather than functioning as attractants by themselves. In contrast, nine-component synthetic HIPV blends formulated in the same ratio and concentration as in a natural sample was as attractive as the original headspace sample. By providing further insights into maize–stemborer–parasitoid tritrophic interactions, results from this study will help in selecting crop cultivars emitting appropriate HIPVs to attract natural enemies and perhaps repel damaging crop pests.

Effects of Abiotic Factors on HIPV-Mediated Interactions between Plants and Parasitoids.

In contrast to constitutively emitted plant volatiles (PV), herbivore-induced plant volatiles (HIPV) are specifically emitted by plants when afflicted with herbivores. HIPV can be perceived by parasitoids and predators which parasitize or prey on the respective herbivores, including parasitic hymenoptera. HIPV act as signals and facilitate host/prey detection. They comprise a blend of compounds: main constituents are terpenoids and “green leaf volatiles.” Constitutive emission of PV is well known to be influenced by abiotic factors like temperature, light intensity, water, and nutrient availability. HIPV share biosynthetic pathways with constitutively emitted PV and might therefore likewise be affected by abiotic conditions. However, the effects of abiotic factors on HIPV-mediated biotic interactions have received only limited attention to date. HIPV being influenced by the plant's growing conditions could have major implications for pest management. Quantitative and qualitative changes in HIPV blends may improve or impair biocontrol. Enhanced emission of HIPV may attract a larger number of natural enemies. Reduced emission rates or altered compositions, however, may render blends imperceptible to parasitoides and predators. Predicting the outcome of these changes is highly important for food production and for ecosystems affected by global climate change.

A tritrophic approach to the preference–performance hypothesis involving an exotic and a native plant

Exotic plants often generate physical and chemical changes in native plant communities where they become established. A major challenge is to understand how novel plants may affect trophic interactions in their new habitats, and how native herbivores and their natural enemies might respond to them. We compared the oviposition preference and offspring performance of the crucifer specialist, Pieris brassicae, on an exotic plant, Bunias orientalis, and on a related native plant, Sinapis arvensis. Additionally, we studied the response of the parasitoid, Cotesia glomerata to herbivore-induced plant volatiles (HIPV) and determined the volatile blend composition to elucidate which compound(s) might be involved in parasitoid attraction. On both host plants we also compared the parasitism rate of P. brassicae by C. glomerata. Female butterflies preferred to oviposit on the native plant and their offspring survival and performance was higher on the native plant compared to the exotic. Although, headspace analysis revealed qualitative and quantitative differences in the volatile blends of both plant species, C. glomerata did not discriminate between the HIPV blends in flight-tent bioassays. Nevertheless, parasitism rate of P. brassicae larvae was higher on the native plant under semi-field conditions. Overall, P. brassicae oviposition preference may be more influenced by bottom-up effects of the host plant on larval performance than by top-down pressure exerted by its parasitoid. The potential for dietary breadth expansion of P. brassicae to include the exotic B. orientalis and the role of top-down processes played by parasitoids in shaping herbivore host shifts are further discussed.

Non‐pathogenic rhizobacteria interfere with the attraction of parasitoids to aphid‐induced plant volatiles via jasmonic acid signalling

Beneficial soil-borne microbes, such as mycorrhizal fungi or rhizobacteria, can affect the interactions of plants with aboveground insects at several trophic levels. While the mechanisms of interactions with herbivorous insects, that is, the second trophic level, are starting to be understood, it remains unknown how plants mediate the interactions between soil microbes and carnivorous insects, that is, the third trophic level. Using Arabidopsis thaliana Col-0 and the aphid Myzus persicae, we evaluate here the underlying mechanisms involved in the plant-mediated interaction between the non-pathogenic rhizobacterium Pseudomonas fluorescens and the parasitoid Diaeretiella rapae, by combining ecological, chemical and molecular approaches. Rhizobacterial colonization modifies the composition of the blend of herbivore-induced plant volatiles. The volatile blend from rhizobacteria-treated aphid-infested plants is less attractive to an aphid parasitoid, in terms of both olfactory preference behaviour and oviposition, than the volatile blend from aphid-infested plants without rhizobacteria. Importantly, the effect of rhizobacteria on both the emission of herbivore-induced volatiles and parasitoid response to aphid-infested plants is lost in an Arabidopsis mutant (aos/dde2-2) that is impaired in jasmonic acid production. By modifying the blend of herbivore-induced plant volatiles that depend on the jasmonic acid-signalling pathway, root-colonizing microbes interfere with the attraction of parasitoids of leaf herbivores.

Function of defensive volatiles in pedunculate oak (Quercus robur) is tricked by the moth Tortrix viridana

The indirect defences of plants are comprised of herbivore-induced plant volatiles (HIPVs) that among other things attract the natural enemies of insects. However, the actual extent of the benefits of HIPV emissions in complex co-evolved plant-herbivore systems is only poorly understood. The observation that a few Quercus robur L. trees constantly tolerated (T-oaks) infestation by a major pest of oaks (Tortrix viridana L.), compared with heavily defoliated trees (susceptible: S-oaks), lead us to a combined biochemical and behavioural study. We used these evidently different phenotypes to analyse whether the resistance of T-oaks to the herbivore was dependent on the amount and scent of HIPVs and/or differences in non-volatile polyphenolic leaf constituents (as quercetin-, kaempferol- and flavonol glycosides). In addition to non-volatile metabolic differences, typically defensive HIPV emissions differed between S-oaks and T-oaks. Female moths were attracted by the blend of HIPVs from S-oaks, showing significantly higher amounts of (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT) and (E)-β-ocimene and avoid T-oaks with relative high fraction of the sesquiterpenes α-farnesene and germacrene D. Hence, the strategy of T-oaks exhibiting directly herbivore-repellent HIPV emissions instead of high emissions of predator-attracting HIPVs of the S-oaks appears to be the better mechanism for avoiding defoliation.

A key volatile infochemical that elicits a strong olfactory response of the predatory mite Neoseiulus californicus, an important natural enemy of the two-spotted spider mite Tetranychus urticae

Herbivore-induced plant volatiles (HIPVs) emitted from lima bean leaves infested with the two-spotted spider mites Tetranychus urticae strongly attract the predatory mites Neoseiulus californicus. Among these HIPVs, methyl salicylate and linalool can attract the predators. Three green-leaf volatiles (GLVs) of (Z)-3-hexen-1-ol, (Z)-3-hexenyl acetate and (E)-2-hexenal, found in the odor blends from T. urticae-infested leaves and physically damaged leaves, can also attract the predators. To search for a strong predator attractant, the olfactory responses of N. californicus to each synthetic compound or their combinations were investigated in a Y-tube olfactometer. When presented a choice between a mixture of the five compounds (i.e. the two HIPVs and the three GLVs) and T. urticae-infested leaves, N. californicus did not discriminate between these odor sources. The same trend was observed when either a mixture of the two HIPVs or methyl salicylate vs. T. urticae-infested leaves were compared. In contrast, the predators preferred T. urticae-infested leaves to linalool, each of the three GLVs, or a mixture of the three GLVs. These results indicated that methyl salicylate is a strong predator attractant, and its potential attractiveness almost equaled that of the blend of HIPVs from T. urticae-infested leaves.

Chemical and Molecular Ecology of Herbivore-Induced Plant Volatiles: Proximate Factors and Their Ultimate Functions

In response to herbivory, plants emit specific blends of herbivore-induced plant volatiles (HIPVs). HIPVs mediate sizable arrays of interactions between plants and arthropods, microorganisms, undamaged neighboring plants or undamaged sites within the plant in various ecosystems. HIPV profiles vary according to the plant and herbivore species, and the developmental stages and conditions of the live plants and herbivores. To understand the regulatory mechanisms underling HIPV biosynthesis, the following issues are reviewed here: (i) herbivore-induced formation of plant volatile terpenoids and green leaf volatiles; (ii) initial activation of plant responses by feeding herbivores; and (iii) the downstream network of the signal transduction. To understand the ecological significance of HIPVs, we also review case studies of insect-plant and inter-/intraplant interactions mediated by HIPVs that have been documented in the field and laboratory in recent years.