Leaf volatiles and polyphenols in pear trees infested byPsylla pyricola. Evidence of simultaneously induced responses

Summary Feeding by the homopteranPsylla pyricola on leaves of pear trees induces the production of volatile compounds, such as (E,E)--farnesene and methyl-salicylate, as well as the production of polyphenols. The inference on induction is based on GC-MS and HPLC chromatograms from the same samples ofPsylla infested leaves, leaves from the same pear tree beforePsylla infestation and uninfested leaves from other pear trees.Psylla infestation greatly enhanced the production of volatiles ((E,E)--farnesene, methyl-salicylate and others) and triggered the production of new polyphenols, characterized by much longer retention times.However, the responses to infestation depend critically on leaf age (defined by leaf distance to apex). With respect to the leaf volatiles it appears that infested, old leaves produce fewer compounds and lower amounts of the volatiles than infested, young leaves. Moreover, there seem to be differences in pattern. Relative to (E,E)--farnesene, methyl-salicylate was found in much lower amounts in heavily infested, old leaves. With respect to polyphenols it was found that infested old leaves collected in August have polyphenols with the same retention times, but more or less equal amounts as uninfested young leaves collected in May. This shows thatPsylla infestation causes the induced response mostly in young leaves.The induced leaf volatiles may act as synomones to heteropteran bugs. As shown elsewhere,Anthocoris nemoralis responds significantly to (E,E)--farnesene and methyl-salicylate when offered in pure form against clean air in a Y-tube olfactometer. The effect of polyphenols on the performance ofP. pyricola is not yet known. Hence, a role in direct defence is still to be investigated.     

New insights in analysing parasitoid attracting synomones: early volatile emission and use of stir bar sorptive extraction

Summary. It is well known that feeding by Pieris brassicae caterpillars on cabbage leaves triggers the release of volatiles that attract natural antagonists such as the parasitoid Cotesia glomerata. The temporal dynamics in the emissions of parasitoid attracting volatiles has never been elucidated in this system. In a time course experiment, caterpillar infested leaves attracted the parasitoid within one hour after infestation. At such an early stage of infestation, as much as fifty percent of the parasitoids flew towards the infested plant in a wind tunnel bioassay, while only five percent flew towards the non-infested control plant. Three hours after infestation and later, the response to the volatiles from the infested plant reached its maximum and then continued at a constantly high level for the remaining 14 hours of the experiment. Chemical analyses of volatiles collected from infested leaves at short time intervals during the first 24 hours identified a total of ten compounds, comprising green leaf volatiles, terpenoids, and a nitrile. Significant increase of emission within the first 5 hours following initial herbivory was detected for (Z)-3-hexen-1-ol, (Z)-3-hexen-1-yl acetate, cineole and benzylcyanide. Subsequently, a coupled bioassay-chemical analysis procedure was developed allowing for testing and analyzing the same sample for future identification of the bioactive compounds. This was achieved by using stir bar sorptive extraction for the analysis of solvent extracts of caterpillar-damaged leaves.     

Aphid alarm pheromone (E)-β-farnesene: A host finding kairomone for the aphid primary parasitoidAphidius uzbekistanicus (Hymenoptera: Aphidiinae)

Summary The present study aimed to test the possible function of the aphid alarm pheromone (E)--farnesene (EBF) as a host finding kairomone for aphid primary parasitoids. Extracts of volatile emissions of undisturbed aphids and of aphids under parasitoid attack were obtained by air entrainment. Extracts of cornicle secretions were gained by disturbing aphids and taking their secretions into solution. Extracts were compared by gas chromatography. Only air entrainments of aphids under attack and solvent extracts of cornicle secretions contained the alarm pheromone. In Y-tube olfactometer bioassays, femaleA. uzbekistanicus were attracted to aphid groups under attack of parasitoids, presumably by released EBF. High concentrations of synthetic EBF (1.4 µg to 5.7 µg) also attractedA. uzbekistanicus females. Females with oviposition experience reacted more readily to lower concentrations of EBF than females without experience. In experiments designed as Petri-dish bioassays, the test animals could contact filter paper discs that were treated with solutions containing EBF. Behavioural effects like antennation or stinging attack were not observed. With computer video analysis of parasitoid movements, some effects onA. uzbekistanicus behaviour were detected, again indicating attraction to EBF.As the volatile aphid alarm pheromone attractedA. uzbekistanicus females, it can be concluded that this chemical stimulus acts as a host finding kairomone for this parasitoid species. However, its effect over long distances seems to be limited due to the relatively high concentrations required for reactions. Of two other parasitoid species examined (P. volucre andL. testaceipes) onlyP. volucre was also significantly attracted to the volatile EBF in the Y-tube olfactometer.     

Potential parasitoid attractants,volatile composition throughout a bark beetle attack

Several hymenopteran parasitoids prey on mature bark beetle larvae (Scolytidae), concealed under the bark of conifers. According to previous results, certain oxygenated monoterpenes are crucial in host location. However, synthetic baits tested in laboratory bioassays are not quite as attractive to the parasitoids as the natural volatile bouquet from conifers containing susceptible bark beetle larvae. The aim of the current study was to identify the "missing components" in previously tested synthetic baits. Volatiles were sampled throughout bark beetle attacks (Ips typographus) on Norway spruce logs (Picea abies), and analyzed (qualitatively and quantitatively) by coupled gas chromatography-mass spectrometry (GC-MS). In addition, a pilot test was done employing the relatively new technique, fast GC-surface acoustic wave detector (SAW), zNose^TM, described in the methods. Quantitative differences in volatile composition were found between infested and uninfested logs, and between different stages of the bark beetle attack. In addition, the volatile composition of logs with or without mature bark beetle larvae (susceptible to parasitism) was compared, isolating odor differences at this most suitable host stage. In general, the quantity of monoterpenes (the predominant phytochemical volatiles in conifers) was found to decrease with time after initial infestation. In contrast, certain oxygenated monoterpenes and benzenoid compounds increased as susceptible bark beetle larvae developed, which therefore could be candidates for improving parasitoid baits. To the best of our knowledge, this is the first report on long-term dynamics of the volatile composition of bark beetle infested P. abies. The results should be interpreted in the light of previous electrophysiological and behavioral investigations of larval parasitoids, and present a number of compounds that could improve synthetic parasitoid baits.     

Local and systemic induced responses to cabbage root fly larvae (Delia radicum) in Brassica nigra and B. oleracea

Summary. Feeding by belowground herbivores may induce systemic changes in shoot defence levels that affect the performance of above ground herbivores and higher trophic levels. In this paper two wild Brassica species, B. nigra and B. oleracea were experimentally infested with 10 larvae of the cabbage root fly, Delia radicum. Plant dry masses and glucosinolate levels in shoots, main roots, and fine roots were determined at 3, 7, 12 and 14 days after infestation and compared to those of control plants. The systemic response in the leaves differed between plant species. In B. nigra shoot glucosinolate levels in D. radicum infested plants steadily increased with time until they were almost twice those of controls 14 days after infestation. B. oleracea plants infested with D. radicum did not show significant changes in shoot glucosinolate levels within 14 days, which may be due to the unexpected poorer performance of D. radicum on this species. Both plant species showed a local increase in indole glucosinolates in the main roots, which are the preferred feeding site of D. radicum larvae. B. oleracea plants however showed a stronger (1.9 – 4.7 times) increase in indole glucosinolate levels than B. nigra (1.5 – 2.6 times). The increase in indole glucosinolates in B. nigra main roots, was counterbalanced by a significant decrease in aromatic glucosinolate levels. These differences in local responses to D. radicum feeding between the two species may have contributed to the slower growth rates of the larvae on B. oleracea. D. radicum feeding did not result in altered glucosinolate levels in the fine roots in either plant species. The differences in glucosinolate induction patterns between the summer annual B. nigra and the perennial B. oleracea are discussed in the light of their different life histories.     

Mother’s choice of the oviposition site: balancing risk of egg parasitism and need of food supply for the progeny with an infochemical shelter?

Summary. Oviposition site selection of herbivorous insects depends primarily on host plant presence which is essential for offspring survival. However, parasitoids can exploit host plant cues for host location. In this study, we hypothesised that herbivores can solve this dilemma by ovipositing within high plant diversity. A diverse plant species composition might represent an ‘infochemical shelter’, as a potentially complex volatile blend can negatively affect the host location ability of parasitoids. We examined this exemplarily for the egg-laying response of the generalist leaf beetle, Galeruca tanaceti, in relation to (1) host plant availability and (2) plant species diversity in the field. Further, we investigated the effect of odours from mixed plant species compositions on (3) leaf beetle oviposition site selection and on (4) the orientation of its specialised egg parasitoid, Oomyzus galerucivorus. In the field, egg clutch occurrence was positively related to the presence and quantity of two major host plants, Achillea millefolium (yarrow) and Centaurea jacea, and to the number of herbaceous plant species. In two-choice bioassays, female beetles oviposited more frequently on sites surrounded by an odour blend from a diverse plant species composition (including yarrow) than on sites with a pure grass odour blend. In the presence of yarrow odour and an odour blend from a diverse plant mixture (including yarrow) no difference in the oviposition response was recorded. Experienced parasitoid females were attracted to yarrow odours, but showed no response when yarrow odours were offered simultaneously with odours of a non-host plant. In conclusion, it could be shown in laboratory bioassays that the parasitoid responds only to pure host plant odours but not to complex odour blends. In contrast, the herbivore prefers to oviposit within diverse vegetation in the field and in the laboratory. However, the laboratory results also point to a priority of host plant availability over the selection of a potential ‘infochemical shelter’ for oviposition due to high plant diversity.     

Emission of oilseed rape volatiles after pollen beetle infestation; behavioural and electrophysiological responses in the parasitoid <Emphasis Type="Italic">Phradis morionellus</Emphasis>

Summary. The pollen beetle, Meligethes aeneus, is an important pest of oilseed rape, Brassica napus. Larvae of this species feed only in the buds and flowers of Brassicaceae. One important natural enemy of this beetle is the parasitoid Phradis morionellus that attacks larvae in buds and flowers and also feeds on the flowers. The preferences for odours of non-infested and infested rape were tested for both starved and fed parasitoids in Y-tube olfactometer experiments. The volatile blend released from pollen beetle-infested and non-infested flowering rape and from pollen beetle larvae was identified and quantified. Gas chromatography-electroantennodetection analyses were performed with female P. morionellus. Parasitoids in both treatment groups preferred infested rape, but the proportion of responding female P. morionellus was significantly lower for the group that was starved. Six of the 20 volatiles identified were released at higher rates from infested rape than from non-infested. None of these compounds was found in pollen beetle larvae headspace. P. morionellus antennae detected both major and minor components in the volatile blend. The volatiles released at a significantly higher rate from infested rape and detected by P. morionellus antennae were (Z)-3-hexenylacetate, (Z)-3-hexenol, 3-butenyl isothiocyanate and (E,E)-α-farnesene.     

Attractive and repellent host odours guide ticks to their respective feeding sites

Summary. We have studied on-host behaviour of adults of the brown ear tick (Rhipicephalus appendiculatus Neumann, 1901) and the red-legged tick (R. evertsi Neumann, 1897), which prefer to feed mainly inside the ears and the anal regions of bovids respectively. Both species were found to be relatively successful in orienting toward and locating their respective feeding sites from different parts of the host body. Our observations suggested the operation of both avoidance (closer to the feeding site of the other) and attraction (closer to its own feeding site) responses of the ticks. In the laboratory, odour trapped from cattle ears attracted R. appendiculatus but repelled R. evertsi, whereas that from the anal region had an opposite effect. This odour-based push-pull pair of stimuli may largely account for efficient orientation behaviour of the two tick species to their respective feeding sites. We propose that such concurrent deployment of repulsive and attractive cues may be quite widespread among arthropods and related organisms that specialise on specific hosts or microenvironments in the performance of their biological functions.     

Diet-induced chemical phytomimesis by twig-like caterpillars of Biston robustum Butler (Lepidoptera: Geometridae)

Summary. Polyphagous caterpillars of the giant geometer Biston robustum resemble the twigs of their respective food sources in color and shape. Common predatory ants, including Lasius and Formica, were often observed to freely prowl directly on caterpillars bodies, even after antennal contact. This suggests that the cuticular chemicals of the caterpillars resemble those of the twigs of the foodplants, so we analyzed both by GC and GC-MS. The chemical compositions differed among caterpillars fed on a cherry, Prunus yedoensis, a chinquapin Castanopsis cuspidata, and a camellia Camellia japonica. The cuticular chemicals of the caterpillars resembled those of their corresponding food sources. When the caterpillar diets were switched from the cherry to camellia or chinquapin at the 4^th instars, the caterpillars cuticular chemicals changed after molting to resemble those of their respective foods. Caterpillars also changed their cuticular chemicals when they perched on cherry twigs and fed on camellia or chinquapin leaves, but not when they perched on camellia or chinquapin twigs and fed on cherry leaves. The chemical similarities between the caterpillars and the twigs were due to the digestion of host leaves, which indicates that this is a diet-induced adaptation.     

Oviposition-stimulating activity of (E)-capsaicin identified in Capsicum annuum fruit and related compounds towards Helicoverpa assulta (Lepidoptera: Noctuidae)

Summary. The oviposition-stimulating activity of (E)-capsaicin identified in the fruits of red pepper, Capsicum annuum L. (Solanaceae), towards the oriental tobacco budworm, Helicoverpa assulta (Guenée), was examined in the laboratory by using choice and no-choice bioassays. Results were compared with those of the commercially available capsanthin and dihydrocapsaicin, another known constituents of C. annuum fruit, and (Z)-capsaicin. In no-choice oviposition bioassays at 10.5 μg/cm², (E)-capsaicin was the most active oviposition stimulant. Weak stimulatory activity was observed with capsanthin, (Z)-capsaicin and dihydrocapsaicin. In choice oviposition bioassays at 2.6 μg/cm², there were significant differences in oviposition-stimulating activity between (E)-capsaicin and each of the other three alkaloids. Structure-activity relationships indicate that the presence of double bond and its trans configuration appeared to play a crucial role in the oviposition-stimulating activity. On the basis of our results, (E)-capsaicin acts as a contact oviposition stimulant of female H. assulta.     

Response of the predatorRhizophagus grandis to host plant derived chemicals inDendroctonus micans larval frass in wind tunnel experiments (Coleoptera: Rhizophagidae,Scolytidae)

Summary Adults of the predatory beetleRhizophagus grandis are strongly attracted to both adult and larval frass of its specific prey,Dendroctonus micans, in walking bioassays. Spruce bark and resin are relatively unattractive. Solvent extracts of larval frass that were attractive toR. grandis adults in a flight wind tunnel contained a mixture of monoterpenes of host plant origin. A synthetic mixture of these monoterpenes, (+)—-pinene, (–)—-pinene, -phellandrene,dl limonene and 3-carene, was responsible for 70–80% of the activity of the most attractive extract. We suggest that a blend of monoterpenes in frass acts as a kairomone forR. grandis but that attraction to monoterpenes is only the first of a sequence of behavioural responses that ensures successful host location and identification.     

<Emphasis Type="Italic">Ceroplastes albolineatus</Emphasis>, the first scale insect shown to sequester pyrrolizidine alkaloids from its host-plant <Emphasis Type="Italic">Pittocaulon praecox</Emphasis>

Summary. The occurrence of pyrrolizidine alkaloids (PAs) in Pittocaulon (ex Senecio) praecox (Asteraceae) a species endemic to Mexico was established. The aboveground plant organs contain the 1,2-saturated monoester 7-angeloyl-dihydroxyheliotridane together with a small proportion of its 9-angeloyl isomer as major alkaloid. The monoesters are accompanied by the macrocyclic otonecine derivative senkirkine. Roots contain only related macrocyclic PAs with senecionine, senkirkine and platyphylline as major components; monoesters are absent. The broom-like succulent stems of P. praecox are infested by the scale insect Ceroplastes albolineatus conspicuously visible by its huge wax cover. All life-history stages, i.e. females, eggs, first instar nymphs (crawlers) and the wax cover were found associated with PAs. The measured PA concentrations clearly indicate sequestration. The highest PA concentrations (mg / g dry weight) reached are: mature females, 0.44; eggs, 0.58; crawlers, 0.37; wax cover, 0.08. The host plant as well as in the infesting scale insect contain the PAs exclusively as free bases. As a phloem-feeder C. albolineatus must acquire the PAs with the ingested phloem sap. This appears plausible since in Senecio species PA are transmitted and circulated through the phloem path. It is suggested that PAs may protect particularly the crawlers as the most endangered stage in the life-cycle of the scale insect.     

The effects of larval experience with a complex plant latex on subsequent feeding and oviposition by the cabbage looper moth: Trichoplusia ni (Lepidoptera: Noctuidae)

Summary. Leaf disc choice and oviposition bioassays were used to examine the effects of larval experience with a Hoodia gordonii latex on subsequent behaviors. The latex deterred feeding and oviposition by “naïve” cabbage looper (Trichoplusia ni, Noctuidae) larvae and moths with no previous exposure to the material. “Experienced” insects, reared on a diet with the H. gordonii latex (1000 ppm), exhibited lesser feeding deterrence relative to naïve insects. Experienced female moths actually preferred to lay eggs on treated rather than control leaves. There was no observed transfer of behavioral preferences from experienced parents to their offspring. Our results suggest that moths may be acquiring oviposition preferences from larval feeding experience as described by Hopkins’ host selection principal (HHSP) or through chemical legacy.     

Furanocoumarins and their detoxification in a tri-trophic interaction

The parsnip webworm, Depressaria pastinacella, specializes on wild parsnip, Pastinaca sativa, and several species of Heracleum, hostplants rich in toxic furanocoumarins. Rates of furanocoumarin metabolism in this species are among the highest known for any insect. Within its native range in Europe, webworms are heavily parasitized by the polyembryonic parasitoid wasp Copidosoma sosares. In this study, we determined whether these parasitoids are exposed to furanocoumarins in host hemolymph, whether they can metabolize furanocoumarins, and whether parasitism influences the ability of webworms to detoxify furanocoumarins. Hemolymph of webworms fed artificial diet containing 0.3 % fresh weight xanthotoxin, a furanocoumarin prevalent in wild parsnip hosts, contained trace amounts of this toxin; as well, hemolymph of webworms consuming P. sativa flowers and fruits contained trace amounts of six of seven furanocoumarins present in the hostplant. Thus, parasitoids likely encounter furanocoumarins in host hemolymph. Assays of xanthotoxin metabolism in C. sosares failed to show any ability to metabolize this compound. Parasitized webworms, collected from populations of Heracleum sphondylium in the Netherlands in 2004, were on average 55 % larger by weight than unparasitized individuals. This weight is inclusive of host and parasitoid masses. Absolute rates of detoxification (nmoles min⁻¹) of five different furanocoumarins were indistinguishable between parasitized and unparasitized ultimate instars, suggesting that the intrinsic rates of metabolism are fixed. Thus, although parasitized larvae are larger, detoxification rates are not commensurate with size; rates in parasitized larvae expressed per gram of larval mass were 25 % lower than in unparasitized larvae.     

Constitutive and herbivore‐induced volatiles in pear,alder and hawthorn trees

Summary. Qualitative and quantitative differences among pear cultivars were found in constitutive and Cacopsylla-induced volatiles, depending on experimental treatment of the trees (i.e., uninfested and partly or completely infested by psyllids). Blend differences were also found between pear cultivars and wild-type pear, alder and hawthorn–the latter trees are frequently present in pear orchard hedgerows. ?Interesting differences were found in the presence of methyl salicylate and (E,E)-α-farnesene, two compounds previously found to mediate attraction of predatory bugs towards psyllid-infested pear trees. Methyl salicylate is expressed constitutively and is induced systemically by infestation in the whole plant of all four cultivars. (E,E)-α-farnesene on the other hand showed also systemic induction in Bartlett, NY10355 and Beurré Hardy, but in partially infested Conference trees it was induced locally, only in herbivore-damaged leaves. No methyl salicylate or (E,E)-α-farnesene were identified in honeydew. In field collected headspace samples of alder leaves infested by aphids and leaf beetles we found methyl salicylate but no (E,E)-α-farnesene, whereas in uninfested hawthorn neither were identified. Insight in the variability of damage-related pear volatiles will have important implications for integrated pest management in the field. Received 27 August 2002; accepted 28 November 2002 R1D=" Correspondence to: Petru Scutareanu, e-mail:scuterea@science.uva.nl     

Variation in composition of predator-attracting allelochemicals emitted by herbivore-infested plants: Relative influence of plant and herbivore

Summary During foraging, natural enemies of herbivores may employ volatile allelochemicals that originate from an interaction of the herbivore and its host plant. The composition of allelochemical blends emitted by herbivore-infested plants is known to be affected by both the herbivore and the plant. Our chemical data add new evidence to the recent notion that the plants are more important than the herbivore in affecting the composition of the volatile blends. Blends emitted by apple leaves infested with spider mites of 2 different species,T. urticae andP. ulmi, differed less in composition (principally quantitative differences for some compounds) than blends emitted by leaves of two apple cultivars infested by the same spider-mite species,T. urticae (many quantitative and a few qualitative differences). Comparison between three plant species — apple, cucumber and Lima bean — reveals even larger differences between volatile blends emitted upon spider-mite damage (many quantitative differences and several qualitative differences).     

Population dynamics of beech scale (Cryptococcus fagisuga) (Coccina,Pseudococcidae) related to physiological defence reactions of attacked beech trees (Fagus sylvatica)

Summary Changes in the density of beech scale infestation (Cryptococcus fagisuga) of a mature beech stand (Fagus sylvatica) were investigated after gap-cutting in limed and unlimed areas bordering on the gaps and in untreated areas over a 5-year period. Parallelly the concentrations of sucrose, protein amino acids, and procyanidins were examined in the inner and outer bark of non-infested beech trees and beeches infested by beech scale.Irrespective of liming in the areas bordering on the gaps the proportion of beech trees with increasing beech scale infestation was significantly higher, the proportion of beeches with diminishing infestation was significantly lower than in the untreated control areas. Trees with the same infestation tendency occurred in clusters.Physiological defence reactions in the inner and outer bark of the infested beech trees were registrated as an increase in the procyanidin content and a decrease in the protein amino acid content. A change in the outer bark—inner bark—ratio of the infested beech trees showed a transfer of compounds between the inner and outer bark. During the period of observation long-lasting shifts occurred in the pattern of compounds of the bark irrespective of actual infestation intensity.     

Faecal shield chemical defence is not important in larvae of the tortoise beetle Chelymorpha reimoseri (Chrysomelidae: Cassidinae: Stolaini)

The most frequently investigated defence mechanism among larvae of tortoise beetles is protection by faecal shields, which generally present chemicals that are directly sequestered and/or modified from larval host-plants. In the present work we investigate the tortoise beetle Chelymorpha reimoseri that feeds on the leaves of Ipomoea carnea fistulosa (Convolvulaceae), seeking for the importance of this mechanism to their larvae. We show that 4th instar larvae suffer low predation regardless of the presence of shields in field and laboratory bioassays with ants and chicks. Chloroform extract from larvae without shields provided high protection against predation as shown in bioassays in the field, as well as against chicks, suggesting that C. reimoseri does not rely on the shield for protection. The aqueous extract of the shields did not show activity in such bioassays. The compounds responsible for this protection have yet to be identified, and it remains an open question as to whether they are sequestered from the host plant or de novo biosynthesized. This is the first record of chemical defence in cassidine beetles without the need for faecal shields. These findings indicate that more attention should be paid to chemicals present in the tissues of larvae and/or adults of tortoise beetles; the protective compounds sequestered from host plants or de novo biosynthesized can provide an alternative or complementary strategy against predation in these insects.     

Secondary metabolites of the leaf surface affected by sulphur fertilisation and perceived by the cabbage root fly

Summary. Surrogate leaves treated with methanolic leaf surface extracts of Brassica napus L. (cv Express) plants that received three different sulphur fertilisation treatments showed even more marked differences by the oviposition choice of Delia radicum L. than the potted plants. This confirms that the oviposition preference of D. radicum is mediated by chemical compounds on the leaf surface and that the quality of host-plants in terms of their nutrition status can be perceived by the female insect.The oviposition data were positively correlated with the content of fractionated surface extracts containing either CIF (cabbage identification factor; 1,2-dehydro-3-thia- 4,10,10b-triaza-cyclopenta[.a.]fluorine-1-carboxylic acid) or glucosinolates. Electrophysiological recordings from the tarsal chemoreceptor sensilla C₅ and D_3,4 showed that receptor neurons react to glucosinolate- and CIF-fractions. We found that the chemosensory activity of specific glucosinolate- and CIF-receptor neurons corresponded with the respective behavioural activity in the oviposition choice assays. The responses of D. radicum to glucosinolates in the electrophysiological recordings studies corresponded to the observed oviposition preference on plants or artificial leaves characterised with an higher amount of glucosinolates on leave surfaces. The presented data suggested that CIF and glucosinolates are involved in host-plant preference of D. radicum and are perceived by tarsal chemoreceptors.     

The aggregation pheromone ofIps duplicatus and its role in competitive interactions withI. typographus (Coleoptera: Scolytidae)

Summary Ips duplicatus withI. typographus co-inhabiting Norway spruce (Picea abies (L.) Karst.) would benefit from a pheromone blend distinct from that of the larger competitorI. typographus. GC-MS analysis showed thatI. duplicatus males feeding in the host produced ipsdienol (Id),cis-verbenol (cV),trans-verbenol (tV), myrtenol (Mt), andE-myrcenol (EM) and traces of 2-methyl-3-buten-2-ol (MB).I. duplicatus produced Id in approximately racemic form (48.9-54.5% (+)-(S)-isomer). The amounts of Id and EM released over a 9 day period had a maximum of 250 and 5 ng/h/male, respectively, on day 2. Exposure ofI. duplicatus males to myrcene and -pinene resulted in the production of small amounts of Id, cV, tV, Mt, andtrans-pinocarveol, but not of EM. In laboratory bioassays with walking beetles, the pheromone component Id alone was weakly attractive while EM was inactive, but in binary combination with Id strongly synergized attraction. A combination of EM and Id at a release rate equivalent to 100–200 males was more attractive in the field than 70 unmated males in a spruce log. The addition of myrcene ( a suggested pheromone precursor of Id) to Id did not enhance trap caches, while addition of EM increased catches > 10-fold. Subtracting EM from a blend of Id, EM, cV and MB drastically reduced trap catches while subtraction of cV or MB or both had no significant effect. Addition of EM over a wide concentration range to the synthetic pheromone ofI. typographus did not reduce the attraction of females of this species in the laboratory. A two-species pheromone interaction field test releasingI. typographus pheromone components (MB + cV) at 10–1000 male equivalents (ME) andI. duplicatus pheromone (Id + EM) at 0, 10–1000 ME in all possible combinations showed both positive intraspecific dose-response effects and an interspecific inhibition. Higher release rates of EM appeared to inhibitI. typographus, especially males. In a tree colonization model, the response of the two competing species to their respective pheromones show a good separation during the mass-attack with a small initial cross-attraction. It remains to be shown whether either of the two pheromone systems have in fact evolved in the present sympatry, or if they are an incidental effect of ancestry of these phylogenetically distantIps.