Effects of Allelochemicals from First (Brassicaceae) and Second (Myzus persicae and Brevicoryne brassicae) Trophic Levels on Adalia bipunctata

Three Brassicaceae species, Brassica napus (low glucosinolate content), Brassica nigra (including sinigrin), and Sinapis alba (including sinalbin) were used as host plants for two aphid species: the generalist Myzus persicae and the specialist Brevicoryne brassicae. Each combination of aphid species and prey host plant was used to feed the polyphagous ladybird beetle, Adalia bipunctata. Experiments with Brassicaceae species including different amounts and kinds of glucosinolates (GLS) showed increased ladybird larval mortality at higher GLS concentrations. When reared on plants with higher GLS concentrations, the specialist aphid, B. brassicae, was found to be more toxic than M. persicae. Identification of GLS and related degradation products, mainly isothiocyanates (ITC), was investigated in the first two trophic levels, plant and aphid species, by high-performance liquid chromatography and gas chromatography–mass spectrometry, respectively. While only GLS were detected in M. persicae on each Brassicaceae species, high amounts of ITC were identified in B. brassicae samples (allyl-ITC and benzyl-ITC from B. nigra and S. alba, respectively) from all host plants. Biological effects of allelochemicals from plants on predators through aphid prey are discussed in relation to aphid species to emphasize the role of the crop plant in integrated pest management in terms of biological control efficacy.     

Olfactory Responses to Aphid and Host Plant Volatile Releases: (E)-β-Farnesene an Effective Kairomone for the Predator Adalia bipunctata

The volatiles released from several aphid and host plant species, alone or associated, were studied for their infochemical role in prey location. Using a four-arm olfactometer, the attraction of several combinations of three aphid (Myzus persicae, Acyrthosiphon pisum, and Brevicoryne brassicae) and three plant (Vicia faba, Brassica napus, and Sinapis alba) species toward Adalia bipunctata larvae and adults was observed. Both predatory larvae and adults were attracted only by A. pisum and M. persicae when they were crushed, whatever the host plant. (E)-beta-farnesene, the aphid alarm pheromone, was the effective kairomone for the ladybird. Plant leaves alone (V. faba, B. napus, and S. alba) or in association with nonstressed whole aphids (the three species) did not have any attraction for the predator. The B. brassicae specialist aphid is the only prey that was not attracted to A. bipunctata larvae and adults, even if they were crushed. Release of B. brassicae molecules similar to the host plant allelochemicals was demonstrated by GC-MS analysis. The lack of behavioral response of the ladybird at short distance toward the cruciferous specialist aphid was related only to the absence of (E)-beta-farnesene in the aphid prey volatile pattern.     

Heterodera schachtii Nematodes Interfere with Aphid-Plant Relations on Brassica oleracea

Aboveground and belowground herbivore species modify plant defense responses differently. Simultaneous attack can lead to non-additive effects on primary and secondary metabolite composition in roots and shoots. We previously found that aphid (Brevicoryne brassicae) population growth on Brassica oleracea was reduced on plants that were infested with nematodes (Heterodera schachtii) prior (4 weeks) to aphid infestation. Here, we examined how infection with root-feeding nematodes affected primary and secondary metabolites in the host plant and whether this could explain the increase in aphid doubling time from 3.8 to 6.7 days. We hypothesized that the effects of herbivores on plant metabolites would depend on the presence of the other herbivore and that nematode-induced changes in primary metabolites would correlate with reduced aphid performance. Total glucosinolate concentration in the leaves was not affected by nematode presence, but the composition of glucosinolates shifted, as gluconapin concentrations were reduced, while gluconapoleiferin concentrations increased in plants exposed to nematodes. Aphid presence increased 4-methoxyglucobrassicin concentrations in leaves, which correlated positively with the number of aphids per plant. Nematodes decreased amino acid and sugar concentrations in the phloem. Aphid population doubling time correlated negatively with amino acids and glucosinolate levels in leaves, whereas these correlations were non-significant when nematodes were present. In conclusion, the effects of an herbivore on plant metabolites were independent of the presence of another herbivore. Nematode presence reduced aphid population growth and disturbed feeding relations between plants and aphids.     

Effect of Soil Sulphur Levels on Feeding Preference of Brevicoryne brassicae on Brussels Sprouts

Two cultivars of Brussels sprouts, Brassica oleracea var. gemmifera were grown in pots treated with nutrient solutions containing five different levels of soil sulfur. Three replicates, each with 10 pots, were infested with the alate form of the aphid, Brevicoryne brassicae. Another three replicates were not infested with aphids. Total apterae aphids were counted on each leaf of the infested plants four weeks later. At the same time all plants were assayed for total glucosinolates and free thiocyanates. The greatest number of aphids per leaf on infested plants was found for the highest levels of sulfur applied. The highest concentrations of total glucosinolates were found in the leaves of uninfested plants that received the highest levels of sulfur. The lowest level of total glucosinolates was found in the leaves of infested plants that received the highest amount of sulfur, and these leaves had the highest level of free thiocyanates. The results suggest that aphids were attracted in greatest numbers to leaves that had the highest synthesis of glucosinolates and that their presence resulted in reduced glucosinolate levels by a combination of a depression in metabolic rate and a stimulation in myrosinase activity.     

Differential Levels of Insect Herbivory in the Field Associated with Genotypic Variation in Glucosinolates in <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis>

Glucosinolates are commonly found in Arabidopsis thaliana and its crucifer relatives, which are known for their role in defense against insect herbivory. In a common garden experiment, we assessed genotypic variation in glucosinolates in A. thaliana and evaluated the association between this chemistry and both plant damage and fitness. Specifically, glucosinolate concentrations were directly associated with damage levels and inversely associated with fitness. These results are contrary to the general expectation that enhanced chemical defense should result in decreased insect herbivory. As the measured insect community in this field trial was dominated by specialist herbivores, this positive relationship between glucosinolates and herbivory agrees with previous observations that glucosinolates (or their hydrolysis products) attract specialist insects. In addition, glucosinolate diversity in this common garden appeared to affect herbivore damage levels. For example, genotypes that contained alkenyl glucosinolates had higher mean damage levels than those that contained hydroxyalkyl glucosinolates. Results suggest that genotypic variation in glucosinolates may be a major factor in determining plant utilization patterns by insect herbivores in the field.     

Chemical Defenses (Glucosinolates) of Native and Invasive Populations of the Range Expanding Invasive Plant Rorippa austriaca

Due to global warming, species are expanding their range to higher latitudes. Some range expanding plants have become invasive in their new range. The Evolution of Increased Competitive Ability (EICA) hypothesis and the Shifting Defense Hypothesis (SDH) predict altered selection on plant defenses in the introduced range of invasive plants due to changes in herbivore pressures and communities. Here, we investigated chemical defenses (glucosinolates) of five native and seven invasive populations of the Eurasian invasive range expanding plant, Rorippa austriaca. Further, we studied feeding preferences of a generalist and a specialist herbivore among the populations. We detected eight glucosinolates in the leaves of R. austriaca. 8-Methylsulfinyloctyl glucosinolate was the most abundant glucosinolate in all plants. There were no overall differences between native and invasive plants in concentrations of glucosinolates. However, concentrations among populations within each range differed significantly. Feeding preference between the populations by a generalist herbivore was negatively correlated with glucosinolate concentrations. Feeding by a specialist did not differ between the populations and was not correlated with glucosinolates. Possibly, local differences in herbivore communities within each range may explain the differences in concentrations of glucosinolates among populations. Little support for the predictions of the EICA hypothesis or the SDH was found for the glucosinolate defenses of the studied native and invasive R. austriaca populations.     

Brassica Plant Responses to Mild Herbivore Stress Elicited by Two Specialist Insects from Different Feeding Guilds

Compensation growth and chemical defense are two components of plant defense strategy against herbivores. In this study, compensation growth and the response of primary and secondary metabolites were investigated in Brassica rapa plants subjected to infestation by two herbivores from contrasting feeding guilds, the phloem-feeding aphid Brevicoryne brassicae and the leaf-feeding caterpillar Pieris brassicae. These specialist herbivores were used at two different densities and allowed to feed for seven days on a young caged leaf. Changes in growth rates were assessed for total leaf area and bulb mass, whereas changes in primary and secondary metabolites were evaluated in young and mature leaves, roots, and bulbs. Mild stress by caterpillars on young plants enhanced mean bulb mass and elicited a contrasting regulation of aliphatic and indolic glucosinolates in the leaves. In contrast, mild stress by aphids enhanced leaf growth and increased glucosinolate concentrations in the bulb, the most important storage organ of B. rapa. A similar mild stress by either herbivore to older plants did not alter plant growth parameters or concentrations of the metabolites analyzed. In conclusion, Brassica plant growth was either maintained or enhanced under mild herbivore stress, and defense patterns differed strongly in response to herbivore type and plant development stage. These results have implications for the understanding of plasticity in plant defenses against herbivores and for the management of Brassica rapa in agroecosystems.     

Different Narrow-Band Light Ranges Alter Plant Secondary Metabolism and Plant Defense Response to Aphids

Light of different wavelengths affects various physiological processes in plants. Short-wavelength radiation (like UV) can activate defense pathways in plants and enhance the biosynthesis of secondary metabolites (such as flavonoids and glucosinolates) responsible for resistance against certain herbivorous insects. The intensity of light-induced, metabolite-based resistance is plant- and insect species-specific and depends on herbivore feeding guild and specialization. In this study, broccoli (Brassica oleracea var. italica) plants were grown for 4 weeks in a climate chamber under conventional fluorescent tubes and were additionally treated with UV-B (310 nm), UV-A (365 or 385 nm), or violet (420 nm) light generated with UV-B tubes or light-emitting diodes (LEDs). The objective was to determine the influence of narrow bandwidths of light (from UV-B to violet) on plant secondary metabolism and on the performance of the cabbage aphid Brevicoryne brassicae (a specialist) and the green peach aphid Myzus persicae (a generalist). Among flavonol glycosides, specific quercetin and kaempferol glycosides increased markedly under UV-B, while among glucosinolates only 4-methoxy-3-indolylmethyl showed a 2-fold increase in plants exposed to UV-B and UV-A. The concentration of 3-indolylmethyl glucosinolate in broccoli plants increased with UV-B treatment. Brevicoryne brassicae adult weights and fecundity were lower on UV-B treated plants compared to UV-A or violet light-treated plants. Adult weights and fecundity of M. persicae were increased under UV-B and UV-A treatments. When specific light wavelengths are used to induce metabolic changes in plants, the specificity of the induced effects on herbivores should be considered.     

Acclimation to Elevated CO2 Increases Constitutive Glucosinolate Levels of Brassica Plants and Affects the Performance of Specialized Herbivores from Contrasting Feeding Guilds

Plants growing under elevated CO₂ concentration may acclimate by modifying chemical traits. Most studies have focused on the effects of environmental change on plant growth and productivity. Potential effects on chemical traits involved in resistance, and the consequences of such effects on plant-insect interactions, have been largely neglected. Here, we evaluated the performance of two Brassica specialist herbivores from contrasting feeding guilds, the leaf-feeding Pieris brassicae and the phloem-feeding Brevicoryne brassicae, in response to potential CO₂-mediated changes in primary and major secondary metabolites (glucosinolates) in Brassica oleracea. Plants were exposed to either ambient (400 ppm) or elevated (800 ppm) CO₂ concentrations for 2, 6, or 10 weeks. Elevated CO₂ did not affect primary metabolites, but significantly increased glucosinolate content. The performance of both herbivores was significantly reduced under elevated CO₂ suggesting that CO₂-mediated increases in constitutive defense chemistry could benefit plants. However, plants with up-regulated defenses could also be subjected to intensified herbivory by some specialized herbivores, due to a chemically-mediated phagostimulatory effect, as documented here for P. brassicae larvae. Our results highlight the importance of understanding acclimation and responses of plants to the predicted increases in atmospheric CO₂ concentrations and the concomitant effects of these responses on the chemically-mediated interactions between plants and specialized herbivores.     

Analysis of EPG Recorded Probing by Cabbage Aphid on Host Plant Parts with Different Glucosinolate Contents

Probing behavior of adult apterous Brevicoryne brassicae (L.) was monitored with the electical penetration graph (EPG) technique on Sinapis alba L. Old, young, and mature leaves, as well as the generally preferred stems were compared with respect to probing activity and glucosinolate content. Compared to leaves, EPGs on stems showed much prolonged initial probes, which more often led to phloem-phase waveforms and successively to sustained phloem ingestion. However, reaching the first phloem phase within a probe was delayed on stems so that the time to the first phloem phase within the experiment was rather similar on the different plant parts. This suggests that superficial (epidermal and mesophyll) factors are more favorable in stems than in leaves but for deeper tissues, i.e., mesophyll and vascular bundle cells, the situation seems reversed, except for the phloem sieve elements, which seemed more or at least equally suitable. 3-Butenyl, p-hydroxybenzyl, and benzyl glucosinolates were found in all parts. It is concluded that further research on distribution and compartmentation of glucosinolates within plant tissues is needed for a better understanding of the correlations found with some B. brassicae probing parameters.     

Glucosinolate Content and Susceptibility for Insect Attack of Three Populations of Sinapis alba

Sinapis alba is less susceptible to damage by insect pests than Brassica napus. We investigated the composition and distribution of glucosinolates in different plant parts in three populations of S. alba; two populations selected for low-seed-glucosinolate content and one commercial cultivar. We have assessed the susceptibility of low-seed-glucosinolate content populations of S. alba to four insect pests, a flea beetle, a pollen beetle, and two species of aphids. Over 90% of the total glucosinolates in the cotyledons of the three populations of S. alba consisted of sinalbin. There was no difference in feeding damage by flea beetles on different S. alba populations at the cotyledon stage, nor was there a difference in sinalbin concentration of cotyledons. Total glucosinolate levels were highest in younger plant tissues. Sinalbin declined as a proportion of total glucosinolate content in later growth stages, especially in the low breeding lines. Reproduction by aphids was the same on all three populations despite differences in sinalbin content of the S. alba leaves at the growth stage tested. The specialist aphid, Brevicoryne brassicae, was found mainly on young tissues, while the generalist aphid, Myzus persicae, was found predominantly on older plant parts. There was no difference in oviposition by pollen beetles between the S. alba populations, despite the fact that on one population total glucosinolate concentration and the proportion of sinalbin found in the buds were much lower than on the other two populations.     

Modifying the Alkylglucosinolate Profile in <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> Alters the Tritrophic Interaction with the Herbivore <Emphasis Type="Italic">Brevicoryne brassicae</Emphasis> and Parasitoid <Emphasis Type="Italic">Diaeretiella rapae</Emphasis>

Arabidopsis thaliana was used as an experimental model plant to investigate a tritrophic interaction between the plant, a specialist aphid herbivore, Brevicoryne brassicae, and its natural enemy, the parasitoid Diaeretiella rapae. The A. thaliana ecotype Col-5 was transformed with a functional 2-oxoglutarate dependent dioxygenase (BniGSL-ALK) that converts 3-methylsulfinylpropylglucosinolate and 4-methylsulfinylbutylglucosinolate to 2-propenylglucosinolate and 3-butenylglucosinolate, respectively. This transformation results in a change in the glucosinolate hydrolysis profile where 3-butenylisothiocyanate, 2-propenylisothiocyanate and 5-vinyloxazolidine-2-thione are produced in contrast to the wild-type plant where 4-methylsulfinylbutylisothiocyanate is the main product. Performance of B. brassicae was affected negatively by transforming Col-5 with BniGSL-ALK in terms of mean relative growth rates. In a series of behavioral bioassays, naïve D. rapae females were able to discriminate between B. brassicae infested and uninfested Col-5 plants transformed with BniGSL-ALK, with parasitoids showing a preference for B. brassicae infested plants. By contrast, naïve D. rapae females were unable to discriminate between aphid infested and uninfested Col-5 plants. Subsequent air entrainments of B. brassicae infested Col-5 plants transformed with BniGSL-ALK further confirmed the presence of 3-butenylisothiocyanate in the headspace. By contrast, no glucosinolate hydrolysis products were recorded from similarly infested Col-5 plants.     

Influence of Increasing Herbivore Pressure on Modification of Glucosinolate Content of Swedes (Brassica napus spp. rapifera)

The effect of increasing herbivore pressure, in the form of larval feeding damage by the turnip root fly, Delia floralis, on the glucosinolate content of swede roots (Brassica napus ssp. rapifera) was investigated. Only one of the 14 root glucosinolates detected, 3-indolyl methyl glucosinolate, rose significantly with increasing levels of insect attack. Although other root glucosinolate concentrations altered following damage, the induced changes were no greater from inoculation with 20 eggs/root than with 5 eggs/root. Swedes roots that had been damaged by D. floralis contained approximately three times the concentration of total indolyl glucosinolates of control roots. This change was strongly influenced by a fourfold increase in the concentration of 1-methoxy-3-indolyl methyl glucosinolate. The total glucosinolate concentration found in swede roots remained unchanged overall as a result of a fall in the concentration of five of the aliphatic glucosinolates, which balanced the rise in aromatic glucosinolates. The relevance of these results to studies of crucifer–insect interactions are discussed.     

Combined Effect of Intercropping and Turnip Root Fly (<Emphasis Type="Italic">Delia floralis</Emphasis>) Larval Feeding on the Glucosinolate Concentrations in Cabbage Roots and Foliage

The effects of plant competition and herbivory on glucosinolate concentrations in cabbage root and foliage were investigated in a cabbage-red clover intercropping system. Cabbage plants were grown under different competitive pressures and with varying degrees of attack by root-feeding Delia floralis larvae. Glucosinolate concentrations in cabbage were affected both by intercropping and by D. floralis density. Glucosinolate concentrations in foliage generally decreased as a response to intercropping, while the responses to insect root damage of individual glucosinolates were weaker. Root glucosinolates responded more strongly to both intercropping and egg density. Total root glucosinolate concentration decreased with clover density, but only at high egg densities. Increased egg density led to opposite reactions by the indole and aliphatic glucosinolates in roots. The responses of individual root glucosinolates to competition and root damage were complex and, on occasion, nonlinear. Reduced concentrations of several glucosinolates and the tendency towards a decrease in total concentration in cabbage foliage caused by intercropping and larval damage suggest that competing plants or plants with root herbivory do not allocate the same resources as unchallenged plants towards sustaining levels of leaf defensive compounds. This could also be true for root glucosinolate concentrations at high egg densities. In addition, the results suggest that changes occurring within a structural group of glucosinolates may be influenced by changes in a single compound, e.g., glucobrassicin (indol-3-ylmethyl) in foliage or sinigrin (2-propenyl) in roots.     

Sex Pheromone of Cabbage Aphid Brevicoryne brassicae: Identification and Field Trapping of Male Aphids and Parasitoids

The sex pheromone of the cabbage aphid, Brevicoryne brassicae, is shown by GC and GC-MS analysis of pheromone entrained from sexual females, and by electrophysiological studies on single cell preparations from male antennae, to comprise (4aS,7S,7aR)-nepetalactone. The compound proved to be attractive in a laboratory bioassay and release of the pheromone from glass vials placed above water traps in crops of autumn brassicaceous crops increased the catch of males of this species. The specialist parasitoid of B. brassicae, Diaeretiella rapae, and the more general aphid parasitoid Praon volucre were found in significantly larger numbers in pheromone traps as compared to the controls, under certain conditions.     

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.     

A Comparison of Semiochemically Mediated Interactions Involving Specialist and Generalist <Emphasis Type="Italic">Brassica</Emphasis>-feeding Aphids and the Braconid Parasitoid <Emphasis Type="Italic">Diaeretiella rapae</Emphasis>

Diaeretiella rapae, a parasitoid that predominately specializes in the parasitism of Brassica-feeding aphids, attacks Lipaphis erysimi, a specialist feeding aphid of the Brassicaceae and other families in the Capparales, at a greater rate than the generalist-feeding aphid, Myzus persicae. In this study, we investigated the orientation behavior of D. rapae to the volatile chemicals produced when these two aphid species feed on turnip (Brassica rapa var rapifera). We showed no significant preference orientation behavior to either aphid/turnip complex over the other. Isothiocyanates are among the compounds emitted by plants of the Brassicaceae in response to insect feeding damage, including by aphids. We assessed parasitoid orientation behavior in response to laboratory-formulated isothiocyanates. We tested two formulations and discovered significant orientation toward 3-butenyl isothiocyanate. We also assessed plant and aphid glucosinolate content, and showed large levels of glucosinolate concentration in L. erysimi, whereas there was little change in plant content in response to aphid feeding. Our results suggest that during the process of host location, similar cues may be utilized for locating L. erysimi and M. persicae, whereas the acceptance of hosts and their suitability may involve aspects of nonvolatile aphid chemistry.     

Differential Effects of Indole and Aliphatic Glucosinolates on Lepidopteran Herbivores

Glucosinolates are a diverse group of defensive secondary metabolites that is characteristic of the Brassicales. Arabidopsis thaliana (L.) Heynh. (Brassicaceae) lines with mutations that greatly reduce abundance of indole glucosinolates (cyp79B2 cyp79B3), aliphatic glucosinolates (myb28 myb29), or both (cyp79B2 cyp79B3 myb28 myb29) make it possible to test the in vivo defensive function of these two major glucosinolate classes. In experiments with Lepidoptera that are not crucifer-feeding specialists, aliphatic and indole glucosinolates had an additive effect on Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) larval growth, whereas Trichoplusia ni (Hübner) (Lepidoptera: Noctuidae) and Manduca sexta (L.) (Lepidoptera: Sphingidae) were affected only by the absence of aliphatic glucosinolates. In the case of two crucifer-feeding specialists, Pieris rapae (L.) (Lepidoptera: Pieridae) and Plutella xylostella (L.) (Lepidoptera: Plutellidae), there were no major changes in larval performance due to decreased aliphatic and/or indole glucosinolate content. Nevertheless, choice tests show that aliphatic and indole glucosinolates act in an additive manner to promote larval feeding of both species and P. rapae oviposition. Together, these results support the hypothesis that a diversity of glucosinolates is required to limit the growth of multiple insect herbivores.     

Attractiveness of Constitutive and Herbivore-Induced Sesquiterpene Blends of Maize to the Parasitic Wasp <Emphasis Type="Italic">Cotesia marginiventris</Emphasis> (Cresson)

Plant volatile compounds induced by herbivore attack have been demonstrated to provide a signal to herbivore enemies such as parasitic wasps that use these volatiles to locate their hosts. However, in addition to herbivore-induced volatiles, plants often release volatiles constitutively. We assessed the interaction between herbivore-induced and constitutively released volatiles of maize in the attraction of the wasp Cotesia marginiventris that parasitizes herbivorous lepidopteran larvae feeding on maize. Experiments were carried out with olfactometers in which the sources of volatiles were transgenic Arabidopsis thaliana plants overexpressing maize sesquiterpene synthases that produce blends of herbivore-induced or constitutive compounds. We found that the constitutive volatiles of maize terpene synthase 8 (TPS8) were attractive to C. marginiventris, just like the herbivore-induced volatiles of TPS10 studied earlier. A mixture of both the TPS8 and TPS10 volatile blends, however, was more effective in parasitoid attraction, indicating that constitutively released sesquiterpenes enhance the attraction of those induced by herbivores. While C. marginiventris did not distinguish among the volatiles of TPS8, TPS10, nor those of another maize sesquiterpene synthase (TPS5), when these blends were combined, their attractiveness to the wasp appeared to increase with the complexity of the blend.     

Does Aphid Infestation Interfere with Indirect Plant Defense against Lepidopteran Caterpillars in Wild Cabbage?

Attraction of parasitoids to plant volatiles induced by multiple herbivory depends on the specific combinations of attacking herbivore species, especially when their feeding modes activate different defense signalling pathways as has been reported for phloem feeding aphids and tissue feeding caterpillars. We studied the effects of pre-infestation with non-host aphids (Brevicoryne brassicae) for two different time periods on the ability of two parasitoid species to discriminate between volatiles emitted by plants infested by host caterpillars alone and those emitted by plants infested with host caterpillars plus aphids. Using plants originating from three chemically distinct wild cabbage (Brassica oleracea) populations, Diadegma semiclausum switched preference for dually infested plants to preference for plants infested with Plutella xylostella hosts alone when the duration of pre-aphid infestation doubled from 7 to 14 days. Microplitis mediator, a parasitoid of Mamestra brassicae caterpillars, preferred dually-infested plants irrespective of aphid-infestation duration. Separation of the volatile blends emitted by plants infested with hosts plus aphids or with hosts only was poor, based on multivariate statistics. However, emission rates of individual compounds were often reduced in plants infested with aphids plus hosts compared to those emitted by plants infested with hosts alone. This effect depended on host caterpillar species and plant population and was little affected by aphid infestation duration. Thus, the interactive effect of aphids and hosts on plant volatile production and parasitoid attraction can be dynamic and parasitoid specific. The characteristics of the multi-component volatile blends that determine parasitoid attraction are too complex to be deduced from simple correlative statistical analyses.