Induction Of Volatile Organic Compounds Research Articles

Induction of volatile organic compounds in chrysanthemum plants following infection by Rhizoctonia solani.

This study investigated the effects of Rhizoctonia solani J.G. Kühn infestation on the volatile organic compound (VOC) emissions and biochemical composition of ten cultivars of chrysanthemum (Chrysanthemum × morifolium /Ramat./ Hemsl.) to bring new insights for future disease management strategies and the development of resistant chrysanthemum cultivars. The chrysanthemum plants were propagated vegetatively and cultivated in a greenhouse under semi-controlled conditions. VOCs emitted by the plants were collected using a specialized system and analyzed by gas chromatography/mass spectrometry. Biochemical analyses of the leaves were performed, including the extraction and quantification of chlorophylls, carotenoids, and phenolic compounds. The emission of VOCs varied among the cultivars, with some cultivars producing a wider range of VOCs compared to others. The analysis of the VOC emissions from control plants revealed differences in both their quality and quantity among the tested cultivars. R. solani infection influenced the VOC emissions, with different cultivars exhibiting varying responses to the infection. Statistical analyses confirmed the significant effects of cultivar, collection time, and their interaction on the VOCs. Correlation analyses revealed positive relationships between certain pairs of VOCs. The results show significant differences in the biochemical composition among the cultivars, with variations in chlorophyll, carotenoids, and phenolic compounds content. Interestingly, R. solani soil and leaf infestation decreased the content of carotenoids in chrysanthemums. Plants subjected to soil infestation were characterized with the highest content of phenolics. This study unveils alterations in the volatile and biochemical responses of chrysanthemum plants to R. solani infestation, which can contribute to the development of strategies for disease management and the improvement of chrysanthemum cultivars with enhanced resistance to R. solani.

Effect of water availability on volatile‐mediated communication between potato plants in response to insect herbivory

Abstract Airborne plant communication is a widespread phenomenon in which volatile organic compounds (VOCs) from damaged plants boost herbivore resistance in neighbouring, undamaged plants. Although this form of plant signalling has been reported in more than 30 plant species, there is still a considerable knowledge gap on how abiotic factors (e.g. water availability) alter its outcomes. We performed a greenhouse experiment to test for communication between potato plants Solanum tuberosum in response to herbivory by the generalist insect Spodoptera exigua and whether communication was affected by water availability. We paired emitter and receiver potato plants, with half of the emitters damaged by S. exigua larvae and half serving as undamaged controls. Both emitter and receiver plants were factorially subjected to one of two watering treatments: high (i.e. well‐watered) vs. low (i.e. reduced watering) availability, thus effectively teasing apart water availability effects on the emission and reception components of signalling. After 4 days of herbivore feeding, we collected emitter VOCs and receivers were subjected to feeding by S. exigua to test for effects of signalling on induced resistance. Herbivory by S. exigua led to increased VOCs emissions as well as changes in VOCs composition in emitter plants. Furthermore, emitters subjected to low water availability exhibited a weaker induction of VOCs in response to herbivory relative to well‐watered emitters. Results from the feeding bioassay indicated that receivers exposed to VOCs from herbivore‐induced emitters showed lower S. exigua damage (i.e. induced resistance) compared to receivers exposed to undamaged emitters. However, we did not observe a significant effect of water availability in either emitters or receivers on plant signalling effects on receiver resistance. Overall, our study contributes to understanding how the abiotic context affects plant communication by providing evidence of water availability effects on the induction of VOCs that act as airborne signals between plants. The observed changes in induced VOCs due to the water treatment had no detectable consequences for plant communication. Accordingly, the induction of key compounds mediating communication was apparently not compromised by our experimental conditions. Read the free Plain Language Summary for this article on the Journal blog.

Induction of volatile organic compounds in Triticum aestivum (wheat) plants following infection by different Rhizoctonia pathogens is species specific

The most popular means of plant protection is the chemical method, but this control is often connected with the need for repeating chemical treatments. Thus, eco-friendly strategies should be developed where, under the European Green Deal, aromatic plants and their repellent properties seem to constitute a good alternative. In earlier studies, we have shown that insect injury, bacteria infestation and pathogen infection induce plant volatile organic compounds (VOCs) emission, which can provide defensive functions to plants.In this study, Triticum aestivum L. (Poaceae) cv. ‘Jenga’ wheat plants were intentionally infected with one of four Rhizoctonia species (R. cerealis, R. solani, R. zeae, and R. oryzae). The soil was inoculated by the pathogens during sowing, whereas shoots were inoculated at stage BBCH 33. In greenhouse experiments, we measured VOCs from wheat 3, 7 and 11 days following stem infestation, or 42 days following soil inoculation of Rhizoctonia spp. VOC emissions were found to be largest on days 7 or 11 post-stem inoculation (>3 days post-stem inoculation >42 days post-soil inoculation). T. aestivum infected by pathogens induced five common green leaf volatiles (GLVs), namely (Z)-3-hexenal = (Z)-3-HAL, (E)-2-hexenal = (E)-2-HAL, (Z)-3-hexen-1-ol = (Z)-3- HOL, (E)-2-hexenol = (E)-2-HOL, (Z)-3-hexen-1-yl acetate = (Z)-3-HAC], six common terpenes (β-pinene = β-PIN, β-myrcene = β-MYR, Z-ocimene = Z-OCI, linalool = LIN, benzyl acetate = BAC, β-caryophyllene = β-CAR), and indole = IND. We found that R. cerealis infested T. aestivum emitted the largest amounts of (Z)-3-HAL and (Z)-3-HAC, while T. aestivum infested by R. solani released the largest amount of LIN (7 or 11 days following stem infestation). VOCs released by the T. aestivum after R. cerealis (AGD I) and R. solani (AG 5) infestations were significantly larger in comparison to R. zeae (WAG-Z) and R. oryzae (WAG-O) for the volatiles (Z)-3-HAL, (E)-2-HAL, (Z)-3-HOL, (E)-2-HOL, (Z)-3-HAC, β-PIN, β-MYR, and LIN. With the exception of (E)-2-HOL, β-MYR, LIN, BAC, β-CAR, the other VOCs were emitted in similar amounts by infected T. aestivum 3 days following stem and soil inoculation. The quantities of induced VOCs were higher at days 7 and 11 than at 3 days post-infection, and greater when T. aestivum was infected with Rhizoctonia on the stem base than through the soil.

Herbivore Gender Effects on Volatile Induction in Aspen and on Olfactory Responses in Leaf Beetles

Hybrid aspen (Populus tremula × tremuloides Michx.) is a fast-growing tree species used for short-rotation forestry in northern latitudes. Aspen species have a rich herbivore fauna, including defoliating leaf beetles that induce emissions of volatile organic compounds (VOCs) when feeding on aspen leaves. We investigated the differential induction of VOCs by male and female Phratora laticollis leaf beetles feeding on hybrid aspen and the differences in the orientation of beetles in response to gender-specific induced VOCs. The hypotheses for the study were (1) the VOCs in the headspace of plants infested with beetles of the two genders individually and in mixed aggregates would vary subtly, and (2) foraging adult beetles would be able to detect differences in VOC blends and use them to fine-tune their orientation choices. In Y-tube bioassays, both females and males preferred VOCs from leaves damaged by one gender (females or males) over undamaged leaves. However, if leaves were damaged by a two-gender population, neither females nor males indicated a preference over volatiles of undamaged leaves. Leaves damaged by both beetle genders simultaneously had significantly increased green leaf volatile (GLV), benzenoid and homoterpene emissions compared to undamaged leaves. Emissions of these compounds possibly indicate higher herbivore pressure and a higher risk of attack by parasitoids and predators and could thus be the cause of the lack of beetle preference. Our findings provide new basic information on gender-based host plant selection by herbivores and may be helpful in the development of sustainable biogenic VOC-based herbivore-control methods for intensive short-rotation hybrid aspen production.

Species- and density-dependent induction of volatile organic compounds by three mite species in cassava and their role in the attraction of a natural enemy.

Upon damage by herbivores, plants induce an array of volatile organic compounds (VOCs) that mediate ecological interactions involving communication with organisms of thesecond and third trophic levels. VOC-mediated tritrophic interactions have largely been studied in various systems, including cassava (Manihot esculenta), but little is known about the chemical nature of herbivore-induced VOCs in this crop and the response they evoke in natural enemies. Several tetranychid and predatory mites are associated with cassava. Here, VOC emissions from uninfested plants and plants infested with 200 or 400 Mononychellus tanajoa, a specialist herbivore on cassava, and the generalists Tetranychus urticae and T. gloveri were measured. Dual-choice experiments were also conducted to assess the preference of inexperienced (reared on prey-infested bean plants) and experienced (adapted on prey-infested cassava plants) predatory mites, Neoseiulus idaeus (Phytoseiidae), between odors of uninfested plants versus odors of plants infested with M. tanajoa, T. urticae or T. gloveri. Two hundred individuals significantly increased the emissions of (Z)-3-hexen-1-ol, (E)-β-ocimene, β-caryophyllene, alloaromadendrene and (E)-geranyl acetone in T. urticae-infested plants, and (E)-β-ocimene and methyl salicylate (MeSA) in T. gloveri-infested plants. Four hundred individuals significantly increased the emissions of (Z)-3-hexen-1-ol, MeSA, α-pinene and D-limonene in M. tanajoa-infested plants. In addition, T. urticae at this density induced (E)-β-ocimene, D-limonene, (E)-geranyl acetone and six compounds that were not detected in other treatments. Tetranychus gloveri-infested plants induced the emissions of (E)-2-hexenal and D-limonene. Regardless of the infesting species, inexperienced N. idaeus did not discriminate between uninfested or infested plants. Upon experience, they discriminated between the odors of uninfested and T. urticae-damaged plants. Our findings reveal that mite infestations in cassava result in density-dependent and species-specific emission of VOCs, and that N. idaeus relies on associative learning to forage for its prey.

Interactive effects of aphid feeding and virus infection on host gene expression and volatile compounds in salt-stressed soybean, Glycine max (L.) Merr.

Saline soils are becoming an important limiting factor in production agriculture. Soybean cultivars [Glycine max (L.) Merr.] differ in their ability to tolerate salt stress with those that cannot limit ion uptake into leaves being salt sensitive. Those that can partially limit ion uptake into leaves are generally more salt tolerant. Soybean mosaic virus (SMV) is an important viral pathogen of soybean worldwide and is commonly transmitted by the soybean aphid, Aphis glycines Matsumura. In this study, we investigate the interaction of salt stress in soybean with SMV infection and infestation by the soybean aphid by measuring aphid populations in a no-choice assay, gene expression levels, and the induction of volatile organic compounds using static headspace GC–MS analysis. Salt stress and SMV infection both reduced total aphid populations, though SMV did not reduce the total number of aphids per gram of fresh weight. Aphid suppression of a calcium EF hand gene and OPR1 was lost when salt-sensitive soybean plants were salt stressed and when salt-tolerant plants were subjected to all three stressors. The relative levels of SMV in aphid-infested soybeans were increased by salt stress in the salt-sensitive cultivar, whereas SMV levels decreased in the salt-tolerant cultivar. Static headspace collection of volatile organic compounds revealed that salt stress and SMV infection had suppressive activities on aphid-induced terpenes. These results suggest that although salt stress has a negative impact on aphid population size, the changes in volatiles and SMV levels could alter the incidence of SMV in salt-stressed fields.

Volatile organic compounds released by maize following herbivory or insect extract application and communication between plants

AbstractTo protect themselves from herbivory, plants have evolved an arsenal of physical and chemical defences and release a variety of volatile organic compounds (VOCs). By releasing these VOCs, a signalling plant can both reduce herbivory, sometimes by more than 90%, and also warn neighbouring plants about an attack. The aim of this study was to assess the influence of herbivory and insect extract application on VOC release by damaged/treated and nearby undamaged/untreated maize plants. We confirmed that European corn borer (Ostrinia nubilalis) larvae attack or larvae extract application induced maize VOC release. Greater amounts of (Z)‐3‐hexenal, (E)‐2‐hexenal, (Z)‐3‐hexen‐1‐ol, (E)‐2‐hexen‐1‐ol, β‐myrcene, (Z)‐3‐hexen‐1‐yl acetate, 1‐hexyl acetate, (Z)‐ocimene, linalool, benzyl acetate, methyl salicylate, indole, methyl anthranilate, geranyl acetate, β‐caryophyllene, (E)‐β‐farnesene and (Z)‐3‐hexenal, (Z)‐3‐hexen‐1‐ol, (Z)‐3‐hexen‐1‐yl acetate, (Z)‐ocimene, linalool, indole, methyl anthranilate, geranyl acetate, β‐caryophyllene and (E)‐β‐farnesene were released as a result of biotic stress after insect attack or insect extract application. The amounts of each VOC released were qualitatively and quantitatively distinct and dependent on time after biotic stress exposure. However, for all biotic stresses, significantly lower VOC induction was measured when leaves were damaged/treated for three days, as compared to seven days. Our work also demonstrated that undamaged/untreated neighbouring plants also release significant amounts of VOCs. This suggests that VOC emission by a damaged/treated plant stimulates VOC induction in nearby undamaged/untreated plants. However, the concentrations of all VOCs released by neighbouring undamaged/untreated maize plants were lower than those from damaged/treated plants and were negatively correlated with distance from a damaged/treated plant. Still, significant VOC induction occurred in undamaged/untreated plants even at 3 m distance from a damaged/infected plant. Our work suggests that maize plant protective defence responses (VOC emission) can be induced via application of European corn borer extracts.

Botrytis cinerea infection in three cultivars of chrysanthemum in ‘Alchimist’ and its mutants: Volatile induction of pathogen-infected plants

Three closely related chrysanthemum cultivars were examined in terms of the effect of Botrytis cinerea infection and leaves injury on the emission rates of fourteen volatile organic compounds (VOCs). Mechanical injury or pathogen infestation of vegetative tissues can induce VOCs production, which can provide defensive functions to plants. Infection with B. cinerea induces the plant defense system in chrysanthemum leaves and stimulates the emission of VOCs that may deter herbivores and attack natural enemies. The change in VOCs emission, as a result of injury or fungal infection in chrysanthemum, has not been studied yet.Our results indicate that a pathogen can induce several VOCs in chrysanthemum plants. In the greenhouse, we measured VOCs from chrysanthemum 3 or 7 days following foliar inoculation. We confirmed that several green leaf volatiles ((GLVs); (Z)-3-hexenal, (E)-2-hexenal, (Z)-3-hexen-1-ol, (E)-2-hexen-1-ol, (Z)-3-hexen-1-yl acetate), terpenes (β-pinene, β-myrcene, (Z)-ocimene, linalool, β-caryophyllene, (E)-β-farnesene), and shikimic acid pathway derivatives (benzyl acetate, methyl salicylate, indole) were positively induced from chrysanthemum plants infected with B. cinerea (Pietrowska et al., 2015). The quantities of the VOCs induced were higher 7 than 3 post infection days. Control plants released only very little but detectable amounts of VOCs. We also set up additional control plants (pierced plants) to facilitate pathogen infection. It was an additional stress and we collected odors also from those plants. In this case the quantities of the VOCs induced were higher 3 than 7 post-injury days. Also our paper suggests a large qualitative and quantitative overlap of VOCs induction from chrysanthemum plants with a mechanical injury when compared to both mechanical injury and the injury from fungal pathogens.

Maize Voc Induction after Infection by the Bacterial Pathogen, Pantoea ananatis, Alters Neighbouring Plant Voc Emissions

Plants induce volatile organic compounds (VOCs) after pathogen infection and exposure to a neighbouring infected plant. In a greenhouse, we measured VOCs from maize cv. ‘Prosna’ 3 or 7 d following foliar, or 42 d following soil, inoculation of Panteoa ananatis, a bacterial pathogen, as well as from uninfected neighbouring maize located 1 or 3 m from each infected plant treatment. We predicted the degree of VOC induction to be greatest 7 d post-foliar > 3 d post-foliar > 42 d post-soil inoculation; also, infected plant VOC induction > 1 m uninfected neighbour > 3 m uninfected neighbour. Maize infected by P. ananatis induced six common green leaf volatiles (GLVs), four common terpenes, and one common skikimic acid pathway derivative. Our results in general confirmed these two predictions, but there was an interaction. While 3 d post-foliar inoculated plants had greater VOC induction than an uninfected neighbour exposed 1 m from a 7 d post-foliar inoculated plant, 42 d post-soil inoculated plants had lower VOC induction than an uninfected neighbour exposed 1 m from a 3 d post-foliar inoculated plant. Thus, infected maize did not always emit higher VOC concentrations than uninfected maize exposed to an infected plant; it depended on the route of infection (foliar vs. soil inoculation) for the infected plant and which infected plant treatment was exposed to an uninfected plant. The VOC blend of maize cv. ‘Prosna’ after P. ananatis infection appears to be quantitatively different when compared to infection by a Fusarium spp. blend. The relevance of these maize VOC blend differences after infection by different pathogens needs to be studied, but our results suggest that maize responds with quantitatively different VOC blends depending on the infecting pathogen.

Volatile organic compounds released by Rumex confertus following Hypera rumicis herbivory and weevil responses to volatiles

AbstractWe report in this study large induction of volatile organic compounds (VOCs) from a single inflorescence of mossy sorrel (Rumex confertus Willd., Polygonaceae), by herbivory of the weevil (Hypera rumicis L., Coleoptera: Curculionidae). VOCs blend induced by the weevil herbivory included 1 green leaf volatiles (GLVs) ((Z)‐3‐hexen‐1‐yl acetate), five terpenes ((Z)‐β‐ocimene, linalool, geranyl acetate, β‐caryophyllene and (E)‐β‐farnesene), three esters (benzyl acetate, methyl salicylate and methyl anthranilate) and one aromatic heterocyclic organic compound (indole). Uninjured plants produced only detectable amounts of VOCs. A Y‐tube experiment revealed that both females and males of H. rumicis were not attracted to any of tested concentrations (1, 5, 25, 125 ng/min). Also both females and males were significantly repelled by the highest concentrations (25 and 125 ng/min). Additionally, concentration of 5 ng/min proved to be repellent for females of H. rumicis.

Synthesis of 6-substituted 1-oxoindanoyl isoleucine conjugates and modeling studies with the COI1-JAZ co-receptor complex of lima bean.

The conjugates of 6-substituted 1-oxoindanoyl carboxylic acids with L-isoleucine are mimics of the plant hormone (+)-7-iso-JA-L-Ile (3) that controls and regulates secondary metabolism and stress responses. In order to generate ligands that can be used as hormone-like compounds possessing different biological activities, an efficient and short synthesis of 6-bromo-1-oxoindane-4-carboxylic acid opens a general route to 6-Br-1-oxoindanoyl L-isoleucine conjugate (Br-In-L-Ile) (9a) as a key intermediate for several bioactive 6-halogen-In-L-Ile analogs (7a, 8a, 10a). The 6-ethynyl-In-L-Ile analog (11a) might be a valuable tool to localize macromolecular receptor molecules by click-chemistry. The activities of In-Ile derivatives were evaluated by assays inducing the release of volatile organic compounds (VOCs) in lima bean (Phaseolus lunatus). Each compound showed slightly different VOC induction patterns. To correlate such differences with structural features, modeling studies of In-Ile derivatives with COI-JAZa/b/c co-receptors of P. lunatus were performed. The modeling profits from the rigid backbone of the 1-oxoindanonoyl conjugates, which allows only well defined interactions with the receptor complex.

Apion miniatum Germ. Herbivory on the Mossy Sorrel, Rumex confertus Willd.: Induced Plant Volatiles and Weevil Orientation Responses

In this paper we report on the large induction of volatile organic compounds (VOCs) from a single leaf of the mossy sorrel, Rumex confertus Willd. (Polygonaceae), by herbivory of the red dock weevil (Apion miniatum Germ., Coleoptera: Apionidae). VOC blends induced by the red dock weevil herbivory included 5 green leaf volatiles (GLVs) ((Z)-3-hexenal, (E)-2-hexenal, (Z)-3-hexenol, (E)-2-hexenol, (Z)-3-hexen-1-yl acetate), two esters (benzyl acetate, methyl salicylate), and four terpenes ((Z)-β-ocimene, linalool, βcaryophyllene, (E)-β-farnesene). Not surprisingly, uninjured plants produced only small amounts of VOCs. A Y-tube experiment revealed that females of A. miniatum were attracted to the blend (blend 1) of (Z)-3-hexenal + (E)-2-hexenal + (Z)-3-hexenol + (E)-2-hexenol + (Z)-3-hexen-1-yl acetate at two concentrations of 1 and 5 ng·min -1 . In contrast males of the red dock weevil were not attracted to any tested concentration. Moreover, females and males of A. miniatum were not attracted to any concentration of blend 2 ((Z)-ocimene + linalool + benzyl acetate + methyl salicylate + s-caryophyllene + (E)-s-farnesene). Yet the red dock weevil females and males were repelled by the highest concentration (125 ng·min -1 ) for both blends.

Induced defenses of Veronica spicata: Variability in herbivore-induced volatile organic compounds

Plants release volatile organic compounds (VOCs) that have many eco-physiological functions. Induction of plant VOCs is known to occur upon herbivory. Herbivore-induced VOCs are involved in the attraction of predators and parasitoids, a phenomenon known as an indirect defense of plants. We measured the VOC profiles of the wild species Veronica spicata with and without larval feeding and oviposition by the specialist butterfly Melitaea cinxia. V. spicata showed great plasticity when deploying indirect defences. The induction of several ubiquitous terpenoids and green leaf volatiles (GLVs) was associated with larval feeding, whereas the increase of two ketones, 6-methyl-5-hepten-2-one and t-geranylacetone and the suppression of GLVs were associated with oviposition by the butterfly.

Pre-exposure to nitric oxide modulates the effect of ozone on oxidative defenses and volatile emissions in lima bean

The roles that ozone and nitric oxide (NO), the chief O3 precursor, play in the antioxidative balance and inducible volatile emissions of lima bean were assessed. Exposure to O3 inhibited APX, CAT, and GR, decreased GSH content and induced emissions of (E)-β-ocimene, limonene, 1,8-cineole, linalool, (E)-4,8-dimethyl-1,3,7-nonatriene (E)-DMNT, 2-butanone and nonanal. O3 did not induce emissions of (E)-β-caryophyllene and appeared to reduce the antioxidative capacity of plants to a greater extent than NO and NO followed by O3 (NO/O3) treatments. There were significant differences in emissions of (E)-β-ocimene and linalool between NO/O3 treated plants and controls, but no differences in antioxidant concentrations. A model to explain the relationships between the ascorbate–glutathione cycle and O3 and NO inducible volatiles was proposed. Our findings suggest that prior exposure to NO modulates the oxidative effect of ozone by the process of cross-tolerance, which might regulate the antioxidative system and induction of volatile organic compounds.

Synthetic Cis-Jasmone Exposure Induces Wheat and Barley Volatiles that Repel the Pest Cereal Leaf Beetle, Oulema melanopus L.

The plant semiochemical cis-jasmone primes/induces plant resistance that deters herbivores and attracts natural enemies. We studied the induction of volatile organic compounds (VOCs) in winter wheat and spring barley after exposure of plants to three synthetic cis-jasmone doses (50μl of 1, 100, and 1 × 10(4)ngμl(-1)) and durations of exposure (1, 3, and 6h). Cereal leaf beetle, Oulema melanopus, adult behavioral responses were examined in a Y-tube olfactometer to cis-jasmone induced plant VOC bouquets and to two synthetic blends of VOCs (3 green leaf volatiles (GLVs); 4 terpenes + indole). In both cereals, eight VOCs [(Z)-3-hexanal, (Z)-3-hexanol, (Z)-3-hexanyl acetate, (Z)-β-ocimene, linalool, β-caryophyllene, (E)-ß-farnesene, and indole] were induced 100- to 1000-fold after cis-jasmone exposure. The degree of induction in both cereals was usually positively and linearly associated with increasing exposure dose and duration. However, VOC emission rate was only ~2-fold greater from plants exposed to the highest vs. lowest cis-jasmone exposure doses (1 × 10(4) difference) or durations (6-fold difference). Male and female O. melanopus were deterred by both cereal VOC bouquets after plant exposure to the high cis-jasmone dose (1 × 10(4)ngμl(-1)), while females were also deterred after plant exposure to the low dose (1ngμl(-1)) but attracted to unexposed plant VOC bouquets. Both O. melanopus sexes were repelled by terpene/indole and GLV blends at two concentrations (25ng · min(-1); 125ng · min(-1)), but attracted to the lowest dose (1ng · min(-1)) of a GLV blend. It is possible that the biologically relevant low cis-jasmone dose has ecological activity and potential for inducing field crop VOCs to deter O. melanopus.

Ecological and phytohormonal aspects of plant volatile emission in response to single and dual infestations with herbivores and phytopathogens

SummaryIn their natural environment, plants are faced with a multitude of attackers, of which insect herbivores and plant pathogens are an important component. In response to these attacks, plants release volatile organic compounds (VOCs), which play an important role in the communication between plants and the associated community members, such as other herbivores, phytopathogens and the natural enemies of herbivores.While numerous studies have focused on either plant–pathogen or plant–insect interactions, less is known when these two sets of interactions co‐occur. Depending on the mode of attack of the pathogen (necrotroph vs. biotroph) or herbivore (chewing vs. piercing‐sucking) they will activate different defence pathways in the plant in which the phytohormones salicylic acid (SA), jasmonic acid (JA) and ethylene (ET) play key roles. As these pathways can crosstalk, a pathogen infection can interfere in a plant's defence response to herbivory, and vice versa.Infestation of a plant with organisms inducingSAsignalling prior to – or simultaneously with – attack by organisms that induce theJApathway often suppressesJAsignalling. However, the impact of this signalling pathway crosstalk onVOCinduction is not clear cut, as there is high variability in the effects on volatile emissions, ranging from suppression to enhanced emission. The effects of the modified volatile blends on the foraging success of carnivorous natural enemies of herbivorous insects have started to be investigated. Foraging success of natural enemies generally withstands this modification of the host‐inducedVOCblend, but the presence or absence of key compounds is an important determinant of the response of certain carnivores.Further studies incorporating plant–insect and plant–pathogen interactions at different levels of biological integration will provide valuable insight in how plants integrate signals from different suites of attacking organisms into an adaptive defence response.

Induced plant defense via volatile production is dependent on rhizobial symbiosis

Nitrogen-fixing rhizobia can substantially influence plant-herbivore interactions by altering plant chemical composition and food quality. However, the effects of rhizobia on plant volatiles, which serve as indirect and direct defenses against arthropod herbivores and as signals in defense-associated plant-plant and within-plant signaling, are still unstudied. We measured the release of jasmonic acid (JA)-induced volatiles of rhizobia-colonized and rhizobia-free lima bean plants (Fabaceae: Phaseolus lunatus L.) and tested effects of their respective bouquets of volatile organic compounds (VOCs) on a specialist insect herbivore (Mexican bean beetle; Coccinellidae: Epilachna varivestis Mulsant) in olfactometer choice trials. In a further experiment, we showed that VOC induction by JA reflects the plant responses to mechanical wounding and insect herbivory. Following induction with JA, rhizobia-colonized plants released significantly higher amounts of the shikimic acid-derived compounds, whereas the emission of compounds produced via the octadecanoid, mevalonate and non-mevalonate pathways was reduced. These changes affected the choice behavior of beetles as the preference of non-induced plants was much more pronounced for plants that were colonized by rhizobia. We showed that indole likely represents the causing agent for the observed repellent effects of jasmonic acid-induced VOCs of rhizobia-colonized lima bean plants. Our study demonstrates a rhizobia-triggered efficacy of induced plant defense via volatiles. Due to these findings, we interpret rhizobia as an integral part of legume defenses against herbivores.

Volatile induction of infected and neighbouring uninfected plants potentially influence attraction/repellence of a cereal herbivore

AbstractPathogen infection can induce plant volatile organic compounds (VOCs). We infected ‘McNeal’ wheat and ‘Harrington’ barley with a Fusarium spp. blend (F. graminearum,F. avenaceum and F. culmorum). Both cereals had the greatest VOC induction 14 days after pathogen innoculation, only slightly lower induction occurred at 7 days, but displayed no induction at 1 days. The induced VOC bouquet for both cereals included six green leaf volatiles (GLVs; e.g. (Z)‐3‐hexenol and (Z)‐3‐hexenyl acetate), four terpenes (linalool, linalool oxide, (Z)‐β‐ocimene and (E)‐β‐caryophyllene) and benzyl acetate. Neighbouring, uninfected individuals of both cereals had significant VOC induction when exposed to an infected, conspecific plant. The temporal pattern and VOC blend were qualitatively similar to infected plants but with quantitative reductions for all induced VOCs. The degree of neighbouring, uninfected plant induction was negatively related to distance from an infected plant. Plant VOC induction in response to pathogen infection potentially influences herbivore attraction or repellency. Y‐tube tests showed that herbivorous female and male Oulema cyanella Voet. (Chrysomelidae: Coleoptera) were significantly attracted to (Z)‐3‐hexenal and (Z)‐3‐hexenyl acetate at 300 and 1500 ng/h but were repelled by both GLVs as well as (Z)‐β‐ocimene and linalool at 7500 ng/h. These O. cyanella behavioural responses were significantly at higher concentrations than those emitted by single plants with pathogen‐induced VOCs, so adults might only be able to respond to a dense group of infected plants. Also, O. cyanella dose responses differ from the previously tested congeneric O. melanopus (cereal leaf beetle), which was attracted to three VOCs induced by Fusarium infection of maize, barley and wheat. Future behavioural tests may indicate whether different herbivore dose responses measured with each VOC singly can help to predict attraction or repellency to injured and uninjured VOC bouquets from different host plant species.

Can forest trees compensate for stress-generated growth losses by induced production of volatile compounds?

Plants produce a variety of volatile organic compounds (VOCs). Under abiotic and biotic stresses, the number and amount of produced compounds can increase. Due to their long life span and large size, trees can produce biogenic VOCs (BVOCs) in much higher amounts than many other plants. It has been suggested that at cellular and tree physiological levels, induced production of VOCs is aimed at improving plant resistance to damage by reactive oxygen species generated by multiple abiotic stresses. In the few reported cases when biosynthesis of plant volatiles is inhibited or enhanced, the observed response to stress can be attributed to plant volatiles. Reported increase, e.g., in photosynthesis has mostly ranged between 5 and 50%. A comprehensive model to explain similar induction of VOCs under multiple biotic stresses is not yet available. As a result of pathogen or herbivore attack on forest trees, the induced production of VOCs is localized to the damage site but systemic induction of emissions has also been detected. These volatiles can affect fungal pathogens and the arrival rate of herbivorous insects on damaged trees, but also act as signalling compounds to maintain the trophic cascades that may improve tree fitness by improved efficiency of herbivore natural enemies. On the forest scale, biotic induction of VOC synthesis and release leads to an amplified flow of BVOCs in atmospheric reactions, which in atmospheres rich in oxides of nitrogen (NOx) results in ozone formation, and in low NOx atmospheres results in oxidation of VOCs, removal in ozone from the troposphere and the resulting formation of biogenic secondary organic aerosol (SOA) particles. I will summarize recent advances in the understanding of stress-induced VOC emissions from trees, with special focus on Populus spp. Particular importance is given to the ecological and atmospheric feedback systems based on BVOCs and biogenic SOA formation.

Gastrophysa polygoni herbivory on Rumex confertus: Single leaf VOC induction and dose dependent herbivore attraction/repellence to individual compounds

We report large induction (>65 fold increases) of volatile organic compounds (VOCs) emitted from a single leaf of the invasive weed mossy sorrel, Rumex confertus Willd. (Polygonaceae), by herbivory of the dock leaf beetle, Gastrophysa polygoni L. ( Coleoptera: Chrysomelidae). The R. confertus VOC blend induced by G. polygoni herbivory included two green leaf volatiles (( Z)-3-hexenal, ( Z)-3-hexen-1-yl acetate) and three terpenes (linalool, ß-caryophyllene, ( E)-ß-farnesene). Uninjured leaves produced small constitutive amounts of the GLVs and barely detectable amounts of the terpenes. A Y-tube olfactometer bioassay revealed that both sexes of adult G. polygoni were attracted to ( Z)-3-hexenal and ( Z)-3-hexen-1-yl acetate at a concentration of 300 ng h −1. No significant G. polygoni attraction or repellence was detected for any VOC at other concentrations (60 and 1500 ng h −1). Yet, G. polygoni males and females were significantly repelled by (or avoided) at the highest test concentration (7500 ng h −1) of both GLVs and ( E)-ß-farnesene. Mated male and female G. polygoni might be attracted to injured R. confertus leaves, but might avoid R. confertus when VOC concentrations (especially the terpene ( E)-ß-farnesene) suggest high overall plant injury from conspecifics, G. viridula, or high infestations of other herbivores that release ( E)-ß-farnesene (e.g., aphids). Tests in the future will need to examine G. polygoni responses to VOCs emitted directly from uninjured (constitutive) and injured (induced) R. confertus, and examine whether R. confertus VOC induction concentrations increase with greater tissue removal on a single leaf and/or the number of leaves with feeding injury.