Defensive role ofAllium sulfur compounds for leek mothAcrolepiopsis assectella Z. (Lepidoptera) against generalist predators

It has been shown previously that sulfur volatiles produced byAllium plants affect the behavior of their specialist phytophages and of their specialist entomophages. The action of these compounds in protecting the leek mothAcrolepiopsis assectella against generalist entomophages was studied in comparison to the proposed original defensive role of these compounds against generalist herbivorous insects. Two ants species,Formica selysi andF. fusca, were used as generalist predators. Six behavioral criteria of the predatory behavior of the ants were studied in presence of the last-instar caterpillars (C₅). C₅ reared on artificial diets with or without leek components were tested, as well as C₅ soaked in frass of leek-reared caterpillars or disulfide solutions. In addition, the response of the ants to pure chemicals found in leek was studied using honey solutions with or without sulfur compounds. The sulfur allelochemicals ofAllium plants have a negative action on predatory ants. Interestingly, the nonvolatile precursors of sulfur volatiles ofAllium plants seem to have a protective role for their phytophagous insects against generalist entomophages.     

Effects of Sulfur-Containing Precursors of Allium Secondary Metabolites on Behavior and Physiology of Diadromus pulchellus, a Specialist Parasitoid of Acrolepiopsis assectella

Diadromus pulchellus, a monophagous parasitoid of the leek moth, Acrolepiopsis assectella, a specialist of plants in the Allium genus, was raised on four series of chrysalids coming from four diets: leek leaves (LL), an artificial diet with (D + L) or without (D – L) leek powder, and an artificial diet plus 1% methyl-cysteinesulfoxide, a precursor of characteristic sulfur volatiles (D + P). Various behavioral and physiological responses by the parent parasitoids and the progeny were studied. No differences were observed in behavior among the four series. The higher precursor concentration observed in LL and in D + P than in D + L and D – L was responsible for a lower emergence ratio and higher egg production in LL and D + P. The consequences of these variations are discussed in relation to narrowness of the tritrophic system. The fact that sulfur volatiles favorable to behavior arise from nonvolatile sulfur precursors, having negative effects on physiology, can also contribute to the maintenance of the system.     

Food experience on the predatory behavior of the ant Myrmica rubra towards a specialist moth,Acrolepiopsis assectella

Entomophagous insects are often repelled by the secondary compounds of the plants eaten by their prey. These compounds, therefore, take on a defensive role for the phytophagous species that sequester them. Given that numerous entomophagous species are capable of learning, the effects on the foraging behavior of a repeated experience were investigated in the predatory ant Myrmica rubra. The sulfur amino acids methyl-cysteine sulfoxide (MCSO) and propyl-cysteine sulfoxide (PCSO) produced by Allium plants were identified in caterpillars of the leek moth Acrolepiopsis assectella. Three behavioral studies were carried out, with or without prior familiarization with caterpillars reared either on leek or on an artificial diet containing no Allium compounds. In choice tests with the two types of caterpillars, unfamiliarized ants displayed a preference for caterpillars reared on the artificial diet, but this preference disappeared or was reversed in both young and old ants after familiarization.     

Herbivore-Induced Volatiles in the Perennial Shrub, <Emphasis Type="Italic">Vaccinium corymbosum</Emphasis>, and Their Role in Inter-branch Signaling

Herbivore feeding activates plant defenses at the site of damage as well as systemically. Systemic defenses can be induced internally by signals transported via phloem or xylem, or externally transmitted by volatiles emitted from the damaged tissues. We investigated the role of herbivore-induced plant volatiles (HIPVs) in activating a defense response between branches in blueberry plants. Blueberries are perennial shrubs that grow by initiating adventitious shoots from a basal crown, which produce new lateral branches. This type of growth constrains vascular connections between shoots and branches within plants. While we found that leaves within a branch were highly connected, vascular connectivity was limited between branches within shoots and absent between branches from different shoots. Larval feeding by gypsy moth, exogenous methyl jasmonate, and mechanical damage differentially induced volatile emissions in blueberry plants, and there was a positive correlation between amount of insect damage and volatile emission rates. Herbivore damage did not affect systemic defense induction when we isolated systemic branches from external exposure to HIPVs. Thus, internal signals were not capable of triggering systemic defenses among branches. However, exposure of branches to HIPVs from an adjacent branch decreased larval consumption by 70% compared to those exposed to volatiles from undamaged branches. This reduction in leaf consumption did not result in decreased volatile emissions, indicating that leaves became more responsive to herbivory (or “primed”) after being exposed to HIPVs. Chemical profiles of leaves damaged by gypsy moth caterpillars, exposed to HIPVs, or non-damaged controls revealed that HIPV-exposed leaves had greater chemical similarities to damaged leaves than to control leaves. Insect-damaged leaves and young HIPV-exposed leaves had higher amounts of endogenous cis-jasmonic acid compared to undamaged and non-exposed leaves, respectively. Our results show that exposure to HIPVs triggered systemic induction of direct defenses against gypsy moth and primed volatile emissions, which can be an indirect defense. Blueberry plants appear to rely on HIPVs as external signals for inter-branch communication.     

Exogenous Application of Jasmonic Acid Induces Volatile Emissions in Rice and Enhances Parasitism of <Emphasis Type="Italic">Nilaparvata lugens</Emphasis> Eggs by theParasitoid <Emphasis Type="Italic">Anagrus nilaparvatae</Emphasis>

Jasmonate signaling pathway plays an important role in induced plant defense against herbivores and pathogens, including the emission of volatiles that serve as attractants for natural enemies of herbivores. We studied the volatiles emitted from rice plants that were wounded and treated with jasmonic acid (JA) and their effects on the host-searching behavior of the rice brown planthopper, Nilaparvata lugens (Stål), and its mymarid egg parasitoid Anagrus nilaparvatae Pang et Wang. Female adults of N. lugens significantly preferred to settle on JA-treated rice plants immediately after release. The parasitoid A. nilaparvatae showed a similar preference and was more attracted to the volatiles emitted from JA-treated rice plants than to volatiles from control plants. This was also evident from greenhouse and field experiments in which parasitism of N. lugens eggs by A. nilaparvatae on plants that were surrounded by JA-treated plants was more than twofold higher than on control plants. Analyses of volatiles collected from rice plants showed that JA treatment dramatically increased the release of volatiles, which included aliphatic aldehydes and alcohols, monoterpenes, sesquiterpenes, methyl salicylate, n-heptadecane, and several as yet unidentified compounds. These results confirm an involvement of the JA pathway in induced defense in rice plants and demonstrate that the egg parasitoid A. nilaparvatae exploits plant-provided cues to locate hosts. We explain the use of induced plant volatiles by the egg parasitoid by a reliable association between planthopper feeding damage and egg presence.     

Herbivore-Induced Volatile Production by Arabidopsis thaliana Leads to Attraction of the Parasitoid Cotesia rubecula: Chemical, Behavioral, and Gene-Expression Analysis

Many plant species defend themselves against herbivorous insects indirectly by producing volatiles in response to herbivory. These volatiles attract carnivorous enemies of the herbivores. Research on the model plant Arabidopsis thaliana (L.) Heynh. has contributed considerably to the unraveling of signal transduction pathways involved in direct plant defense mechanisms against pathogens. Here, we demonstrate that Arabidopsis is also a good candidate for studying signal transduction pathways involved in indirect defense mechanisms by showing that: (1) Adult females of Cotesia rubecula, a specialist parasitic wasp of Pieris rapae caterpillars, are attracted to P. rapae-infested Arabidopsis plants. (2) Arabidopsis infested by P. rapae emits volatiles from several major biosynthetic pathways, including terpenoids and green leaf volatiles. The blends from herbivore-infested and artificially damaged plants are similar. However, differences can be found with respect to a few components of the blend, such as two nitriles and the monoterpene myrcene, that were produced exclusively by caterpillar-infested plants, and methyl salicylate, that was produced in larger amounts by caterpillar-infested plants. (3) Genes from major biosynthetic pathways involved in volatile production are induced by caterpillar feeding. These include AtTPS10, encoding a terpene synthase involved in myrcene production, AtPAL1, encoding phenylalanine ammonia-lyase involved in methyl salicylate production, and AtLOX2 and AtHPL, encoding lipoxygenase and hydroperoxide lyase, respectively, both involved in the production of green leaf volatiles. AtAOS, encoding allene oxide synthase, involved in the production of jasmonic acid, also was induced by herbivory.     

Methyl Salicylate,a Soybean Aphid-Induced Plant Volatile Attractive to the Predator <Emphasis Type="Italic">Coccinella septempunctata</Emphasis>

Induced volatiles provide a signal to foraging predatory insects about the location of their prey. In Iowa, early in the growing season of soybean, Glycine max, many predacious seven-spotted lady beetles, Coccinella septempunctata, were observed on plants with heavy infestations of soybean aphid, Aphis glycines. We studied whether the attraction of this beetle is caused by the release of specific volatile compounds of soybean plants infested by aphids. Volatile compounds emitted by soybean plants infested by aphids were compared with those of undamaged, uninfested, and artificially damaged plants. Gas chromatography–mass spectrometry analyses revealed consistent differences in the profiles of volatile compounds between aphid-infested soybean plants and undamaged ones. Significantly more methyl salicylate was released from infested plants at both the V1 and V2 plant growth stages. However, release patterns of two other induced plant volatiles, (d)-limonene and (E,E)-α-farnesene, differed between the two plant growth stages. Gas chromatographic–electroantennographic detection of volatile extracts from infested soybean plants showed that methyl salicylate elicited significant electrophysiological responses in C. septempunctata. In field tests, traps baited with methyl salicylate were highly attractive to adult C. septempunctata, whereas 2-phenylethanol was most attractive to the lacewing Chrysoperla carnea and syrphid flies. Another common lady beetle, the multicolored Asian lady beetle, Harmonia axyridis, showed no preference for the compounds. These results indicate that C. septempunctata may use methyl salicylate as the olfactory cue for prey location. We also tested the attractiveness of some selected soybean volatiles to alate soybean aphids in the field, and results showed that traps baited with benzaldehyde caught significantly higher numbers of aphids.     

Plant-natural enemy association in tritrophic system,Cotesia rubecula-Pieris rapae-brassicaceae (Cruciferae). III: Collection and identification of plant and frass volatiles

To elucidate the identity of the volatile compounds that could be involved in the searching behavior of the parasitoidCotesia rubecula Marshall (Hymenoptera: Braconidae), the volatiles released by cabbage and frass of Lepidoptera feeding on cabbage were collected and analyzed using a gas chromatograph-mass spectrometer. The volatiles emitted by intact cabbage were -pinene, -pinene, myrcene, 1,8-cineole,n-hexyl acetate,cis-3-hexen-1-yl acetate, and dimethyl trisulfide. Mechanical damage on an intact plant induced the release of two more compounds,trans-2-hexenal and 1-methoxy-3-methylene-2-pentanone. Current feeding by larvae ofPieris rapae L. (Pieridae) induced the plant to release all the compounds released after mechanical damage and additionally 4-methyl-3-pentenal and allyl isothiocyanate. Current feeding by larvae ofPlutella xylostella L. (Plutellidae) induced the plant to release all the compounds present after mechanical damage and additionally allyl isothiocyanate. The volatiles emitted after feeding by the lepidopterans had ceased were the same as those emitted by cabbage damaged by mechanical means. The blend of volatiles emitted by frass was comprised of plant chemicals, mainly sulfur compounds. Frass ofP. rapae emitted allyl isothiocyanate, methyl isothiocyanate, methyl propyl sulfide, dimethyl trisulfide,S-methyl methane thiosulfinate, 4-methyl-3-pentenal,trans-2-hexenal, and 2,3-dihydro-4-methyl furan. Frass ofP. xylostella emitted only dimethyl trisulfide andS-methyl methane thiosulfinate. The blend of volatiles emitted by frass is herbivore-species specific.     

Attraction of the Parasitoid <Emphasis Type="Italic">Anagrus nilaparvatae</Emphasis> to Rice Volatiles Induced by the Rice Brown Planthopper <Emphasis Type="Italic">Nilaparvata lugens</Emphasis>

Anagrus nilaparvatae, an egg parasitoid of the rice brown planthopper Nilaparvata lugens, was attracted to volatiles released from N. lugens-infested plants, whereas there was no attraction to volatiles from undamaged plants, artificially damaged plants, or volatiles from N. lugens nymphs, female adults, eggs, honeydew, and exuvia. There was no difference in attractiveness between plants infested by N. lugens nymphs or those infested by gravid females. Attraction was correlated with time after infestation and host density; attraction was only evident between 6 and 24 hr after infestation by 10 adult females per plant, but not before or after. Similarly, after 24 hr of infestation, wasps were attracted to plants with 10 to 20 female planthoppers, but not to plants with lower or higher numbers of female planthoppers. The attractive time periods and densities may be correlated with the survival chances of the wasps' offspring, which do not survive if the plants die before the wasps emerge. Wasps were also attracted to undamaged mature leaves of a rice plant when one of the other mature leaves had been infested by 10 N. lugens for 1 d, implying that the volatile cues involved in host location by the parasitoid are systemically released. Collection and analyses of volatiles revealed that 1 d of N. lugens infestation did not result in the emission of new compounds or an increase in the total amount of volatiles, but rather the proportions among the compounds in the blend were altered. The total amounts and proportions of the chemicals were also affected by infestation duration. These changes in volatile profiles might provide the wasps with specific information on host habitat quality and thus could explain the observed behavioral responses of the parasitoid.     

Systemic release of herbivore-induced plant volatiles by turnips infested by concealed root-feeding larvae Delia radicum L

When attacked by herbivorous insects, many plants emit volatile compounds that are used as cues by predators and parasitoids foraging for prey or hosts. While such interactions have been demonstrated in several host–plant complexes, in most studies, the herbivores involved are leaf-feeding arthropods. We studied the long-range plant volatiles involved in host location in a system based on a very different interaction since the herbivore is a fly whose larvae feed on the roots of cole plants in the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae). The parasitoid studied is Trybliographa rapae Westwood (Hymenoptera: Figitidae), a specialist larval endoparasitoid of D. radicum. Using a four-arm olfactometer, the attraction of naive T. rapae females toward uninfested and infested turnip plants was investigated. T. rapae females were not attracted to volatiles emanating from uninfested plants, whether presented as whole plants, roots, or leaves. In contrast, they were highly attracted to volatiles emitted by roots infested with D. radicum larvae, by undamaged parts of infested roots, and by undamaged leaves of infested plants. The production of parasitoid-attracting volatiles appeared to be systemic in this particular tritrophic system. The possible factors triggering this volatile emission were also investigated. Volatiles from leaves of water-stressed plants and artificially damaged plants were not attractive to T. rapae females, while volatiles emitted by leaves of artificially damaged plants treated with crushed D. radicum larvae were highly attractive. However, T. rapae females were not attracted to volatiles emitted by artificially damaged plants treated only with crushed salivary glands from D. radicum larvae. These results demonstrate the systemic production of herbivore-induced volatiles in this host-plant complex. Although the emission of parasitoid attracting volatiles is induced by factors present in the herbivorous host, their exact origin remains unclear. The probable nature of the volatiles involved and the possible origin of the elicitor of volatiles release are discussed.     

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

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

Herbivore-induced Plant Volatiles Trigger Sporulation in Entomopathogenic Fungi: The Case of <Emphasis Type="Italic">Neozygites tanajoae</Emphasis> Infecting the Cassava Green Mite

A large body of evidence shows that plants release volatile chemicals upon attack by herbivores. These volatiles influence the performance of natural enemies. Nearly all the evidence on the effect of plant volatiles on natural enemies of herbivores concerns predators, parasitoids, and entomophagous nematodes. However, other entomopathogens, such as fungi, have not been studied yet for the way they exploit the chemical information that the plant conveys on the presence of herbivores. We tested the hypothesis that volatiles emanating from cassava plants infested by green mites (Mononychellus tanajoa) trigger sporulation in three isolates of the acaropathogenic fungus Neozygites tanajoae. Tests were conducted under climatic conditions optimal to fungal conidiation, such that the influence of the plant volatiles could only alter the quantity of conidia produced. For two isolates (Altal.brz and Colal.brz), it was found that, compared with clean air, the presence of volatiles from clean, excised leaf discs suppressed conidia production. This suppressive effect disappeared in the presence of herbivore-damaged leaves for the isolate Colal.brz. For the third isolate, no significant effects were observed. Another experiment differing mainly in the amount of volatiles showed that two isolates produced more conidia when exposed to herbivore-damaged leaves compared with clean air. Taken together, the results show that volatiles from clean plants suppress conidiation, whereas herbivore-induced plant volatiles promote conidiation of N. tanajoae. These opposing effects suggest that the entomopathogenic fungus tunes the release of spores to herbivore-induced plant signals indicating the presence of hosts.     

Semiochemical-Mediated Location of Host Habitat by Apanteles carpatus (Say) (Hymenoptera: Braconidae), a Parasitoid of Clothes Moth Larvae

In Y-tube olfactometer bioassays, adult Apanteles carpatus (Say), were attracted to beaver or rabbit pelts infested with larvae of the casemaking clothes moth (CCM)Tinea pellionella L. Porapak Q-captured volatiles from a CCM-infested beaver pelt were also very attractive, whereas isolated CCM larvae or larval feces were not. Coupled gas chromatographic–electroantennographic detection (GC-EAD) analysis of the Porapak Q volatile extract revealed two compounds that elicited responses by A. carpatus antennae. Coupled GC–mass spectrometry (MS) in electron impact and chemical ionization modes of these compounds indicated, and GC-MS and GC-EAD of authentic standards confirmed, that they were nonanal and geranylacetone. While each compound singly did not attract A. carpatusa 1:1 blend of both compounds was as attractive as the volatile extract. Because these compounds are host habitat-derivedA. carpatus must be a habitat rather than host specialist, responding to kairomonal indicators of localized and specific habitats such as animal hair or feather. The tritrophic interaction between A. carpatusits clothes moth hosts and their animal-derived habitats is similar to the well-studied relationship between parasitoids of insect herbivores and their host plant habitats.     

Pollinator and Herbivore Attraction to <Emphasis Type="Italic">Cucurbita</Emphasis> Floral Volatiles

Mutualists and antagonists may place conflicting selection pressures on plant traits. For example, the evolution of floral traits is typically studied in the context of attracting pollinators, but traits may incur fitness costs if they are also attractive to antagonists. Striped cucumber beetles (Acalymma vittatum) feed on cucurbits and are attracted to several volatiles emitted by Cucurbita blossoms. However, the effect of these volatiles on pollinator attraction is unknown. Our goal was to determine whether pollinators were attracted to the same or different floral volatiles as herbivorous cucumber beetles. We tested three volatiles previously found to attract cucumber beetles in a factorial design to determine attraction of squash bees (Peponapis pruinosa), the specialist pollinators of cucurbita species, as well as the specialist herbivore A. vittatum. We found that 1,2,4-trimethoxybenzene was attractive to both the pollinator and the herbivore, indole was attractive only to the herbivore, and (E)-cinnamaldehyde was attractive only to the pollinator. There were no interactions among volatiles on attraction of squash bees or cucumber beetles. Our results suggest that reduced indole emission could benefit plants by reducing herbivore attraction without loss of pollination, and that 1,2,4-trimethoxybenzene might be under conflicting selection pressure from mutualists and antagonists. By examining the attraction of both mutualists and antagonists to Cucurbita floral volatiles, we have demonstrated the potential for some compounds to influence only one type of interaction, while others may affect both interactions and possibly result in tradeoffs. These results shed light on the potential evolution of fragrance in native Cucurbita, and may have consequences for yield in agricultural settings.     

Leek odor analysis by gas chromatography and identification of the most active substance for the leek moth,Acrolepiopsis assectella

When crushed, the leek,Allium porrum emits propyl propanethiosulfinate. The unstable thiosulfinate decomposes during GC analysis in numerous compounds, except on very short columns. The propyl propanethiosulfmate is the most attractive substance for the leek moth,Acrolepiopsis assectella. This thiosulfinate is not active onPlutella xylostella orPlodia interpunctella and seems to be repulsive onEphestia kuehniella.     

The Role of Fresh <Emphasis Type="Italic">versus</Emphasis> Old Leaf Damage in the Attraction of Parasitic Wasps to Herbivore-Induced Maize Volatiles

The odor produced by a plant under herbivore attack is often used by parasitic wasps to locate hosts. Any type of surface damage commonly causes plant leaves to release so-called green leaf volatiles, whereas blends of inducible compounds are more specific for herbivore attack and can vary considerably among plant genotypes. We compared the responses of naïve and experienced parasitoids of the species Cotesia marginiventris and Microplitis rufiventris to volatiles from maize leaves with fresh damage (mainly green leaf volatiles) vs. old damage (mainly terpenoids) in a six-arm olfactometer. These braconid wasps are both solitary endoparasitoids of lepidopteran larvae, but differ in geographical origin and host range. In choice experiments with odor blends from maize plants with fresh damage vs. blends from plants with old damage, inexperienced C. marginiventris showed a preference for the volatiles from freshly damaged leaves. No such preference was observed for inexperienced M. rufiventris. After an oviposition experience in hosts feeding on maize plants, C. marginiventris females were more attracted by a mixture of volatiles from fresh and old damage. Apparently, C. marginiventris has an innate preference for the odor of freshly damaged leaves, and this preference shifts in favor of a blend containing a mixture of green leaf volatiles plus terpenoids, after experiencing the latter blend in association with hosts. M. rufiventris responded poorly after experience and preferred fresh damage odors. Possibly, after associative learning, this species uses cues that are more directly related with the host presence, such as volatiles from host feces, which were not present in the odor sources offered in the olfactometer. The results demonstrate the complexity of the use of plant volatiles by parasitoids and show that different parasitoid species have evolved different strategies to exploit these signals.     

Volatile Emissions from an Odorous Plant in Response to Herbivory and Methyl Jasmonate Exposure

Induced volatile terpenes have been commonly reported among diverse agricultural plant species, but less commonly investigated in odorous plant species. Odorous plants synthesize and constitutively store relatively large amounts of volatiles, and these may play a role in defense against herbivores. We examined the effect of herbivory and methyl jasmonate (MeJA) exposure on the release of volatile organic compounds (VOCs) in the marsh elder, Iva frutescens, which contains numerous constitutive VOCs, mainly mono- and sesquiterpenes. Our specific goal was to test for the presence of inducible VOCs in a naturally occurring plant already armed with VOCs. The abundant, native specialist leaf beetle Paria aterrima was used in herbivore induction trials. VOCs were sampled from herbivore wounded and unwounded, and from MeJA treated and untreated I. frutescens. Total VOC emissions were significantly greater in response to herbivory and MeJA treatment compared to unwounded controls. Herbivore wounding caused a substantial shift in the emission profile (42 VOCs from wounded, compared to 8 VOCs from unwounded I. frutescens), and MeJA had a similar yet less substantial influence on the emission pattern (28 VOCs from MeJA treated compared to 8 VOCs from untreated I. frutescens). Constitutive VOC emissions predominated, but some VOCs were detected only in response to herbivory and MeJA treatment, suggesting de novo synthesis. Several VOCs exhibited a delayed emission profile in contrast to the rapid release of constitutive VOCs, and principal components analysis revealed they were not associated with constitutive emissions. While I. frutescens contains many constitutive VOCs that are released immediately in response to herbivory, it also produces novel VOCs in response to feeding by the specialist P. aterrima and MeJA treatment.     

Variation in Herbivore and Methyl Jasmonate-Induced Volatiles Among Genetic Lines of <Emphasis Type="Italic">Datura wrightii</Emphasis>

Many plant species produce volatile organic compounds after being damaged by herbivores. The production of volatiles also may be induced by exposing plants to the plant hormone, jasmonic acid, or its volatile ester, methyl jasmonate. This study addresses the induction of the production volatile organic compounds among genetic lines of Datura wrightii. Within populations, some plants produce glandular trichomes, whereas others produce nonglandular trichomes, and trichome phenotype is controlled by a single dominant gene. Glandular trichomes not only confer resistance to some herbivorous insects, but they also inhibit many natural enemies of those herbivores. Because of the potential benefit of natural enemies that use volatile cues to find individuals of the non-glandular phenotype, it is reasonable to ask if plants of D. wrightii that differ in trichome morphology might produce different blends of volatile compounds. Volatile compounds were collected from eight genetic lines of plants that had been backcrossed for three generations. Volatiles were collected from pairs of sibling plants before and after insect damage or treatment with methyl jasmonate. Within each pair, one sib expressed glandular trichomes and the other expressed nonglandular trichomes. Overall, plants produced an array of at least 17 compounds, most of which were sesquiterpenes. Total production of volatiles increased from 3.9- to 16.2-fold among genetic lines after insect damage and from 3.6- to 32-fold in plants treated with methyl jasmonate. The most abundant compound was (E)-β-caryophyllene. This single compound comprised from 17 to 59% of the volatiles from insect-damaged plants and from 24 to 88% of the volatiles from plants treated with methyl jasmonate, depending upon genetic line. The production of (E)-β-caryophyllene by the original male parents of the eight genetic lines was significantly related to the mean production of their third-generation backcross progeny indicating that the variation in the production of (E)-β-caryophyllene was inherited. Blends did not differ qualitatively or quantitatively between sibs expressing glandular or nonglandular trichomes.     

Herbivore-Induced Plant Volatiles Mediate In-Flight Host Discrimination by Parasitoids

Herbivore feeding induces plants to emit volatiles that are detectable and reliable cues for foraging parasitoids, which allows them to perform oriented host searching. We investigated whether these plant volatiles play a role in avoiding parasitoid competition by discriminating parasitized from unparasitized hosts in flight. In a wind tunnel set-up, we used mechanically damaged plants treated with regurgitant containing elicitors to simulate and standardize herbivore feeding. The solitary parasitoid Cotesia rubecula discriminated among volatile blends from Brussels sprouts plants treated with regurgitant of unparasitized Pieris rapae or P. brassicae caterpillars over blends emitted by plants treated with regurgitant of parasitized caterpillars. The gregarious Cotesia glomerata discriminated between volatiles induced by regurgitant from parasitized and unparasitized caterpillars of its major host species, P. brassicae. Gas chromatography-mass spectrometry analysis of headspace odors revealed that cabbage plants treated with regurgitant of parasitized P. brassicae caterpillars emitted lower amounts of volatiles than plants treated with unparasitized caterpillars. We demonstrate (1) that parasitoids can detect, in flight, whether their hosts contain competitors, and (2) that plants reduce the production of specific herbivore-induced volatiles after a successful recruitment of their bodyguards. As the induced volatiles bear biosynthetic and ecological costs to plants, downregulation of their production has adaptive value. These findings add a new level of intricacy to plant–parasitoid interactions.     

Role of the Lipoxygenase/lyase Pathway of Host-food Plants in the Host Searching Behavior of Two Parasitoid Species, Cotesia glomerata and Cotesia plutellae

To elucidate the role of the plant lipoxygenase (LOX)/lyase pathway for host search behavior of two parasitic wasps attacking herbivorous larvae, an Arabidopsis mutant (all84) was isolated with a mutation somewhere in the LOX/lyase pathway. Detached leaves of the mutant were shown to release less (Z)-3-hexenal, a first green leaf volatile (GLV) product of the LOX/lyase pathway. The braconid larval parasitoids studied, Cotesia glomerata and Cotesia plutella, differ in their ability to discriminate among plant volatiles induced by feeding of lepidopteran hosts and nonhosts: C. plutella only responds to plant volatiles induced by hosts (Plutella larvae), whereas the response by the more generalist C. glomerata is not host specific. The Arabidopsis mutant all84 infested by Pieris larvae was less attractive to C. glomerata than Arabidopsis wild type (wt) infested by the host larvae. C. glomerata was attracted by two of the GLV biosynthesized through the LOX/lyase pathway, (E)-2-hexenal and (Z)-3-hexenyl acetate. However, attraction of C. plutellae to volatiles from Plutella-infested all84 plants did not differ from attraction to host-infested wt Arabidopsis. Both wasp species were arrested to the respective host-infested edge of the wt leaf by showing characteristic antennal searching behavior on the edge. In C. glomerata, the duration of this searching behavior at the infested leaf edge was significantly shorter on all84 plants than on wt plants. By contrast, the duration of the searching behavior of C. plutellae on the host-infested leaf edge of all84 was not significantly different from that on the wt leaf. These data suggest that the LOX/lyase pathway is directly involved in the production of attractants and arrestants important for host search behavior of the more generalist C. glomerata, but not for the specialist C. plutellae.