Volatiles released from cotton plants in response to<Emphasis Type="Italic"> Helicoverpa zea</Emphasis> feeding damage on cotton flower buds

Feeding of Helicoverpa zea larvae on cotton (Gossypium hirsutum L.) flower buds (squares) for 24 or 48 h induced the release of a number of terpenes [(E)--ocimene, linalool, (E)--farnesene, (E,E)--farnesene, (E)-4,8-dimethyl-1,3,7-nonatriene, (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene], isomeric hexenyl butyrates, 2-methylbutyrates, indole and (Z)-3-hexenyl acetate. These compounds are not released in significant amounts from undamaged squares and freshly damaged squares. The release of inducible compounds was not limited to the damaged squares themselves. The compounds were also released systemically from the upper undamaged leaves of the same plant after 72 h. However, the composition of the blend of systemically released volatiles differed from the blend released by damaged squares. The compounds that were systemically released from undamaged leaves in response to feeding on the squares were (Z)-3-hexenyl acetate, (E)--ocimene, linalool, (E)-4,8-dimethyl-1,3,7-nonatriene, (E)--farnesene, (E,E)--farnesene, and indole. This study shows that insect damage inflicted to the reproductive parts of a plant causes a systemic emission of volatiles from its vegetative parts.     

Aphid antixenosis in cotton is activated by the natural plant defence elicitor cis-jasmone

Upon insect herbivory, plants can release blends of volatile organic compounds (VOCs) that modify herbivore and natural enemy behaviour. We have shown recently that cotton, Gossypium hirsutum, emits a blend of defence VOCs that repels the cotton aphid, Aphis gossypii, upon herbivory by this notorious crop pest, including (Z)-3-hexenyl acetate, (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), methyl salicylate and (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene (TMTT). In this study, we investigated changes in the defence VOC profile of G. hirsutum induced by the naturally-occurring plant elicitor cis-jasmone (CJ) and whether these changes modify the behaviour of A. gossypii. In four-arm olfactometer assays, VOCs from untreated plants were significantly attractive (P < 0.05), whilst VOCs from CJ-treated plants were significantly repellent (P < 0.05). The VOCs induced by CJ appeared to comprise (Z)-3-hexenyl acetate, DMNT, methyl salicylate and TMTT. In quantitative VOC collection studies, sustained release of DMNT and TMTT was observed in CJ-treated plants over a period of five days, with levels becoming statistically significantly higher than for control treated plants on the fifth day in most cases. Despite earlier indications, no statistically significant differences were observed in levels of (Z)-3-hexenyl acetate or methyl salicylate between CJ and control treatments on any day. Furthermore, DMNT and TMTT emissions from CJ-treated plants were further enhanced by subsequent addition of A. gossypii. CJ treatment induced statistically significantly higher DMNT and TMTT expression levels as early as day three, when A. gossypii was present. The results in this study show that CJ can induce the production of A. gossypii-induced VOCs from G. hirsutum, with potential for deployment in novel crop protection strategies.     

Volatiles released from bean plants in response to agromyzid flies

Liriomyza sativae Blanchard and Liriomyza huidobrensis (Blanchard) (Diptera: Agromyzidae) are two invasive flies in China that have caused economical damage on vegetables and ornamental plants. In this article, we report the profiles of emitted volatiles from healthy, mechanically damaged, and leafminer-damaged bean, Phaseolus vulgaris L., plants. Among 25 emitted volatiles identified, (E)-2-hexen-1-al, (3E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), (Z)-3-hexenyl acetate, (Z)-3-hexen-1-ol, (syn)- and (anti)-2-methylpropanal oxime, (syn)-2-methylbutanal oxime, linalool, and (E,E)-α-farnesene were consistently released from damaged bean plants. Combined amounts of these nine compounds made up more than 70% of the total volatiles emitted from each treatment. No qualitative differences in volatile emission were found between bean plants damaged by the two fly species; however, amounts of several major compounds induced by L. huidobrensis damage were significantly higher than those from plants damaged by L. sativae. The mechanically damaged plants released a higher proportion of green leaf volatiles than plants in the other treatments, whereas leafminer-damaged plants produced more terpenoids and oximes. Furthermore, the volatile profiles emitted from plants, damaged by adult leafminers, by second instar larvae, and even the plants with empty mines left by leafminer larvae (the pupal stage) were significantly different. The identification of volatile oximes released from damaged plants was confirmed and is discussed in a behavioral and biological control context.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Do Insectivorous Birds use Volatile Organic Compounds from Plants as Olfactory Foraging Cues? Three Experimental Tests

Some insectivorous birds orient towards insect‐defoliated trees even when they do not see the foliar damage or the herbivores. There are, however, only a few studies that have examined the mechanisms behind this foraging behaviour. Previous studies suggest that birds can use olfactory foraging cues (e.g. volatile organic compounds (VOCs) emitted by defoliated plants), indirect visual cues or a combination of the two sensory cues. VOCs from insect‐defoliated plants are known to attract natural enemies of herbivores, and researchers have hypothesized that VOCs could also act as olfactory foraging cues for birds. We conducted three experiments across a range of spatial scales to test this hypothesis. In each experiment, birds were presented with olfactory cues and their behavioural responses or foraging outcomes were observed. In the first experiment, two different VOC blends, designed to simulate the volatile emissions of mountain birch (Betula pubescens ssp. czerepanovii) after defoliation by autumnal moth (Epirrita autumnata) larvae, were used in behavioural experiments in aviaries with pied flycatchers (Ficedula hypoleuca). The second experiment was a field‐based trial of bird foraging efficiency; the same VOC blends were applied to mountain birches, silver birches (B. pendula) and European white birches (B. pubescens) with plasticine larvae attached to the trees to serve as artificial prey for birds and provide a means to monitor predation rate. In the third experiment, the attractiveness of silver birch saplings defoliated by autumnal moth larvae versus intact controls was tested with great tits (Parus major) and blue tits (Cyanistes caeruleus) in an aviary. Birds did not orient towards either artificial or real trees with VOC supplements or towards herbivore‐damaged saplings when these saplings and undamaged alternatives were hidden from view. These findings do not support the hypothesis that olfactory foraging cues are necessary in the attraction of birds to herbivore‐damaged trees.     

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 the second 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.     

Terpenoids dominate the bouquet of volatile organic compounds produced by <Emphasis Type="Italic">Passiflora edulis</Emphasis> in response to herbivory by <Emphasis Type="Italic">Heliconius erato phyllis</Emphasis> (Lepidoptera: Nymphalidae)

In response to injury, plants produce volatile organic compounds (VOCs) that usually differ depending on the type of damage they have suffered (e.g., mechanical damage, herbivory, and oviposition). The objectives of this study were to identify and compare the bouquet of volatiles emitted by passion vine plants (Passiflora edulis) after injury caused by mechanical damage (MD), herbivory (HB), and oviposition (OV) by the lepidopteran, Heliconius erato phyllis. Following injury, extracts of plant emissions were collected from each treatment every 24 h for three days and were analyzed by GC and GC/MS. Results show that plants emitted 12 volatiles before and after damage, namely terpenoids, ketones, and aldehydes. Although no significant differences were detected between the three treatments individually, if the entire bouquet of volatiles is analyzed, samples collected at 24 h were different from samples collected at 48 and 72 h. However, terpenoid emission increased significantly in HB plants after 24 h. HB plants emitted approximately 6300, 50, 46, 11, 6, and 3.6 times more (3E,7E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene, (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), (E)-β-ocimene, (Z)-β-farnesene, (E)-β-caryophyllene, and farnesane, respectively, compared to control plants. OV plants displayed a peak of emission of (E)-β-ocimene after 72 h, which distinguished them from HB plants. MD plants showed a general increase of VOCs versus undamaged control plants. Furthermore, it has been suggested that (E)-β-ocimene may be sequestered by larvae of H. erato phyllis as a component of the odoriferous bouquet of the abdominal scent glands present in adult males, which play a role in sexual communication.     

Demonstration and characterization of (E)-nerolidol synthase from maize: a herbivore-inducible terpene synthase participating in (3E)-4,8-dimethyl-1,3,7-nonatriene biosynthesis

Upon herbivore attack, maize (Zea mays L.) emits a mixture of volatile compounds that attracts herbivore enemies to the plant. One of the major components of this mixture is an unusual acyclic C₁₁ homoterpene, (3E )-4,8-dimethyl-1,3,7-nonatriene (DMNT), which is also emitted by many other species following herbivore damage. Biosynthesis of DMNT has been previously shown to proceed via the sesquiterpene alcohol, (E )-nerolidol. Here we demonstrate an enzyme activity that converts farnesyl diphosphate, the universal precursor of sesquiterpenes, to (3S)-(E )-nerolidol in cell-free extracts of maize leaves that had been fed upon by Spodoptera littoralis. The properties of this (E )-nerolidol synthase resemble those of other terpene synthases. Evidence for its participation in DMNT biosynthesis includes the direct incorporation of deuterium-labeled (E )-nerolidol into DMNT and the close correlation between increases in (E )-nerolidol synthase activity and DMNT emission after herbivore damage. Since farnesyl diphosphate has many other metabolic fates, (E )-nerolidol synthase may represent the first committed step of DMNT biosynthesis in maize. However, the formation of this unusual acyclic terpenoid appears to be regulated at both the level of (E )-nerolidol synthase and at later steps in the pathway. Received: 20 August 1999 / Accepted: 27 October 1999     

Fusarium infection in maize: volatile induction of infected and neighboring uninfected plants has the potential to attract a pest cereal leaf beetle, Oulema melanopus

Fusarium infection of maize leaves and/or roots through the soil can stimulate the emission of volatile organic compounds (VOCs). It is also well known that VOC emission from maize plants can repel or attract pests. In our experiments, we studied VOC induction responses of Zea mays L. ssp. mays cv. ‘Prosna’ having Fusarium infection (mix of four species) in leaves or roots, then tested for VOC induction of uninfected neighboring plants, and finally examined wind-tunnel behavioral responses of the adult cereal leaf beetle, Oulema melanopus L. (Chrysomelidae: Coleoptera) behavior to four induced VOCs. In the first part of our experiment, 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, 1-hexyl acetate), terpenes (β-pinene, β-myrcene, Z-ocimene, linalool, β-caryophyllene), and shikimic acid pathway derivatives (benzyl acetate, methyl salicylate, indole) were positively induced from maize plants infected by Fusarium spp. The quantities of induced VOCs were higher at 7 d than 3 d post-infection and greater when plants were infected with Fusarium on leaves rather than through soil. In the second part of our experiment, uninfected maize plants also showed significantly positive induction of several VOCs when neighboring an infected plant where the degree of induction was negatively related to the distance from the infected plant. In the third part of our experiment, a Y-tube bioassay was used to evaluate upwind orientation of adult cereal leaf beetles to four individual VOCs. Female and male O. melanopus were significantly attracted to the GLVs (Z)-3-hexenal and (Z)-3-hexenyl acetate, and the terpenes linalool and β-caryophyllene. Our results indicate that a pathogen can induce several VOCs in maize plants that also induce VOCs in neighboring uninfected plants, though VOC induction could increase the range at which an insect pest species is attracted to VOC inducing plants.     

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.     

Priming of Production in Maize of Volatile Organic Defence Compounds by the Natural Plant Activator cis-Jasmone

cis-Jasmone (CJ) is a natural plant product that activates defence against herbivores in model and crop plants. In this study, we investigated whether CJ could prime defence in maize, Zea mays, against the leafhopper, Cicadulina storeyi, responsible for the transmission of maize streak virus (MSV). Priming occurs when a pre-treatment, in this case CJ, increases the potency and speed of a defence response upon subsequent attack on the plant. Here, we tested insect responses to plant volatile organic compounds (VOCs) using a Y-tube olfactometer bioassay. Our initial experiments showed that, in this system, there was no significant response of the herbivore to CJ itself and no difference in response to VOCs collected from unexposed plants compared to CJ exposed plants, both without insects. VOCs were then collected from C. storeyi-infested maize seedlings with and without CJ pre-treatment. The bioassay revealed a significant preference by this pest for VOCs from infested seedlings without the CJ pre-treatment. A timed series of VOC collections and bioassays showed that the effect was strongest in the first 22 h of insect infestation, i.e. before the insects had themselves induced a change in VOC emission. Chemical analysis showed that treatment of maize seedlings with CJ, followed by exposure to C. storeyi, led to a significant increase in emission of the defensive sesquiterpenes (E)-(1R,9S)-caryophyllene, (E)-α-bergamotene, (E)-β-farnesene and (E)-4,8-dimethyl-1,3,7-nonatriene, known to act as herbivore repellents. The chemical analysis explains the behavioural effects observed in the olfactometer, as the CJ treatment caused plants to emit a blend of VOCs comprising more of the repellent components in the first 22 h of insect infestation than control plants. The speed and potency of VOC emission was increased by the CJ pre-treatment. This is the first indication that CJ can prime plants for enhanced production of defensive VOCs antagonist towards herbivores.     

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

Pathogen 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.     

Feeding by emerald ash borer larvae induces systemic changes in black ash foliar chemistry

The exotic wood-boring pest, emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), has been threatening North American ash (Fraxinus spp.) resources, this being recognized since its first detection in Michigan, USA and Ontario, Canada in 2002. Ash trees are killed by larval feeding in the cambial region, which results in disruption of photosynthate and nutrient translocation. In this study, changes in volatile and non-volatile foliar phytochemicals of potted 2-yr-old black ash, Fraxinus nigra Marshall, seedlings were observed in response to EAB larval feeding in the main stem. EAB larval feeding affected levels of six compounds [hexanal, (E)-2-hexenal, (Z)-3-hexenyl acetate, (E)-β-ocimene, methyl salicylate, and (Z,E)-α-farnesene] with patterns of interaction depending upon compounds of interest and time of observation. Increased methyl salicylate emission suggests similarity in responses induced by EAB larval feeding and other phloem-feeding herbivores. Overall, EAB larval feeding suppressed (Z)-3-hexenyl acetate emission, elevated (E)-β-ocimene emission in the first 30 days, but emissions leveled off thereafter, and generally increased the emission of (Z,E)-α-farnesene. Levels of carbohydrates and phenolics increased overall, while levels of proteins and most amino acids decreased in response to larval feeding. Twenty-three amino acids were consistently detected in the foliage of black ash. The three most abundant amino acids were aspartic acid, glutamic acid, glutamine, while the four least abundant were α-aminobutyric acid, β-aminoisobutyric acid, methionine, and sarcosine. Most (16) foliar free amino acids and 6 of the 9 detected essential amino acids decreased with EAB larval feeding. The ecological consequences of these dynamic phytochemical changes on herbivores harbored by ash trees and potential natural enemies of these herbivores are discussed.     

Genetic variation in plant volatile emission does not result in differential attraction of natural enemies in the field

Volatile organic chemical (VOC) emission by plants may serve as an adaptive plant defense by attracting the natural enemies of herbivores. For plant VOC emission to evolve as an adaptive defense, plants must show genetic variability for the trait. To date, such variability has been investigated primarily in agricultural systems, yet relatively little is known about genetic variation in VOCs emitted by natural populations of native plants. Here, we investigate intraspecific variation in constitutive and herbivore-induced plant VOC emission using the native common milkweed plant (Asclepias syriaca) and its monarch caterpillar herbivore (Danaus plexippus) in complementary field and common garden greenhouse experiments. In addition, we used a common garden field experiment to gauge natural enemy attraction to milkweed VOCs induced by monarch damage. We found evidence of genetic variation in the total constitutive and induced concentrations of VOCs and the composition of VOC blends emitted by milkweed plants. However, all milkweed genotypes responded similarly to induction by monarchs in terms of their relative change in VOC concentration and blend. Natural enemies attacked decoy caterpillars more frequently on damaged than on undamaged milkweed, and natural enemy visitation was associated with higher total VOC concentrations and with VOC blend. Thus, we present evidence that induced VOCs emitted by milkweed may function as a defense against herbivores. However, plant genotypes were equally attractive to natural enemies. Although milkweed genotypes diverge phenotypically in their VOC concentrations and blends, they converge into similar phenotypes with regard to magnitude of induction and enemy attraction.     

The social wasp Polybia fastidiosuscula Saussure (Hymenoptera: Vespidae) uses herbivore‐induced maize plant volatiles to locate its prey

Social wasps in the Polybia genus are important for use as pest‐control agents in agricultural systems. The objective of this study was to investigate the behavioural responses of Polybia fastidiosuscula Saussure (Hymenoptera: Vespidae) to volatiles from maize, both constitutive volatiles and those induced by the herbivory of Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae). To assess the behavioural response of P. fastidiosuscula to S. frugiperda larvae, undamaged plants, S. frugiperda‐damaged plants, mechanically damaged plants, mechanically damaged plants plus regurgitant from larvae and extracts from various treatments, bioassays were conducted in a Y‐olfactometer. In addition, the volatiles from plants subjected to different treatments were collected via aeration, and they were quantified and identified. The wasps showed a greater preference for plants with damage induced either by larval feeding or by being mechanically damaged plus regurgitant than for undamaged plants or either larvae alone or mechanically damaged plants. Wasps were more attracted to extracts from plants + S. frugiperda larvae and to an extract from mechanically damaged plants + the regurgitant of larvae compared to hexane. The primary compounds induced by herbivory for 5–6 h after the beginning of the damage or regurgitant treatment were identified as α‐pinene, β‐myrcene, (Z)‐3‐hexenyl acetate, limonene, (E)‐ocimene, linalool, DMNT, (E)‐β‐farnesene, TMTT and indole. The results presented here show that the social wasp P. fastidiosuscula uses herbivore‐induced plant volatiles from maize to locate its prey.     

Timing of induced volatile emissions in maize seedlings

Maize (Zea mays L.) releases specific volatiles in response to herbivory by caterpillars. These volatiles are known to serve as cues for parasitic wasps to locate the herbivores. In the present study the exact time of volatile emission after simulated herbivory (mechanical damage and treatment with caterpillar regurgitant) was measured for seedlings of the cultivars “Ioana Sweet Corn” and “LG11”. Odours were collected every 0.5 h for a total of 12 h. Typical “green leaf odours”, (Z)-3-hexenal, (E )-2-hexenal, (Z)-hexen-1-o1, and (Z)-3-hexen-1-yl acetate, were emitted immediately upon damage and their amounts dropped rapidly after the first collections. Several of the induced compounds were released within 2 h after treatment, while others (mainly sesquiterpenoids) started to be released after 4 h. The LG11 seedlings emitted several compounds (e.g. β-myrcene, (Z)-β-ocimene, benzyl acetate, β-caryophyllene, (E,E )-α-farnesene) that were not detected for Ioana. (E,E )-α-farnesene was continuously emitted by LG11 seedlings, even by undamaged plants. Timing of the release of volatile compounds that the two varieties had in common did not differ significantly, with the exception of indole for which the peak production was considerably earlier for LG11. These findings are discussed in the context of biosynthetic pathways and mechanisms involved in induced emissions of plant volatiles and the exploitation of the resulting odour by parasitoids and predators of herbivores. Received: 23 October 1997 / Accepted: 9 June 1998     

Conversion of airborne nerolidol to DMNT emission requires additional signals in Achyranthes bidentata

DMNT biosynthesis was proposed to proceed via (E)-nerolidol in plants a decade ago. However, (E)-nerolidol function as airborne signal/substrate for in-vivo biosynthesis of DMNT remains to be investigated and the regulation of DMNT production and emission is largely unknown. We address both of these aspects using Achyranthes bidentata model plant in conjunction with deuterium-labeled d(5)-(E)-nerolidol, headspace, GC-FID, and GC/MS-based absolute quantification approaches. We demonstrate that airborne (E)-nerolidol is specifically metabolized in-vivo into DMNT emission, but requires airborne VOC MeJA or predator herbivore as additional environmental signal. In addition, we provide new insight into the complex regulation underlying DMNT emission, and highlight the importance of studying multiple environmental factors on emission patterns of plant VOCs and their mechanistic regulation.     

Induced resistance in mountain birch: defence against leaf-chewing insect guild and herbivore competition

Summary The effect of leaf damage simulating the feeding of early season insect herbivore species, e.g. Epirrita autumnata, to mountain birch, Betula pubescens ssp. tortuosa, on the performance of insect larvae was studied with eleven leaf-chewing sawfly species. I found variation in the results that was due to short- and long-term inducible responses and to the phenology of herbivore species. In general, early and mid-season species were more strongly affected by induced reactions than late-season species. This finding is in accordance with earlier results but I could show that the persistance of induced reactions rather than the influence of timing of damage is responsible for the result. The growth of the larvae of mid-season sawfly species was affected by both short- and long-term induced reactions. This result shows that early season species may escape short-term induced reactions of mountain birch in current year but may not avoid long-term effects. It is supposed that seasonal deterioration of leaf quality either masks the effects of induced defences or late-season species are better adapted to low-quality leaves. Some species show variation in their response to induced defence in different years. This may be due to yearly differences in induced reactions as well as to species-specific responses. Induced defence reactions may play a role in competitive interactions between herbivore species in leaf-chewing guild of mountain birch.     

Systemic induction of volatile release in cotton: How specific is the signal to herbivory?

Plants attacked by herbivorous insects release chemical signals that attract natural enemies of the herbivores to the damaged plants. Feeding of Spodoptera exigua larvae on the lower leaves of cotton (Gossypium hirsutum L.) for multiple feeding periods of 9–12 h with a 12 h, interval in between when the caterpillars are removed overnight, will induce a systemic release of volatile compounds that is comparable to the volatiles released in response to continuous feeding damage on the lower leaves for several days. The systemic volatile release in response to herbivory can be mimicked by mechanically damaging the lower leaves and applying caterpillar oral secretion to the injured leaves over 4 days. Cotton plants that are only mechanically damaged systemically release significantly less -pinene, myrcene, (Z)-3-hexenyl acetate, (E)--farnesene and (E,E)--farnesene after 4 days compared to plants damaged mechanically with application of caterpillar regurgitant. However, multiple 9–12 h mechanical damage alone induces a significantly higher systemic release of (Z)-3-hexenyl acetate, myrcene, (E)--ocimene, and (E)--farnesene after 4 days compared to undamaged control plants. This indicates that multiple mechanical damage alone cannot mimic completely the response induced by mechanically injuring the leaves and applying caterpillar regurgitant. A specific elicitor in the regurgitant of the caterpillar enhances the amount of several systemically released volatiles. Thus, the systemic release of volatile compounds by herbivore-damaged cotton plants appears to be regulated by at least two different mechanisms.     

Effect of mechanical damage on emission of volatile organic compounds from plant leaves and implications for evaluation of host plant specificity of prospective biological control agents of weeds

Assessment of host plant specificity is a critical step in the evaluation of classical biological control agents of weeds which is necessary for avoiding possible damage to non-target plants. Volatile organic compounds (VOCs) emitted by plants likely play an important role in determining which plants attract and are accepted by a prospective arthropod agent. However, current methods to evaluate host plant specificity usually rely on empirical choice and no-choice behavioural experiments, with little knowledge about what chemical or physical attributes are stimulating the insect. We conducted experiments to measure the quantitative and qualitative effects on emission of VOCs caused by simple mechanical damage to leaves of plants known to differ in suitability and attractiveness to a prospective agent. More VOCs were detected from damaged than from undamaged leaves for all three species tested. Discriminant analysis was able to correctly distinguish the taxonomic identity of all plants based on their VOC profiles; however, the VOCs that discriminated species among undamaged leaves were completely different from those that discriminated among damaged leaves. Thus, damaged and undamaged plants present different VOC profiles to insects, which should be considered when conducting host plant specificity experiments. An unacceptable non-target plant, Centaurea cineraria, emitted all except one of the VOCs that were emitted by its preferred host plant, Centaurea solstitialis, indicating the importance of compounds that are repellant in host plant specificity. Centaurea cyanus emitted fewer VOCs than C. solstitialis, which suggests that it lacked some VOCs important for host plant recognition.