Development of a synthetic plant volatile-based attracticide for female noctuid moths. I. Potential sources of volatiles attractive to Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

This paper is the first of a series which will describe the development of a synthetic plant volatile-based attracticide for noctuid moths. It discusses potential sources of volatiles attractive to the cotton bollworm, Helicoverpa armigera (Hübner), and an approach to the combination of these volatiles in synthetic blends. We screened a number of known host and non-host (for larval development) plants for attractiveness to unmated male and female moths of this species, using a two-choice olfactometer system. Out of 38 plants tested, 33 were significantly attractive to both sexes. There was a strong correlation between attractiveness of plants to males and females. The Australian natives, Angophora floribunda and several Eucalyptu s species were the most attractive plants. These plants have not been recorded either as larval or oviposition hosts of Helicoverpa spp., suggesting that attraction in the olfactometer might have been as nectar foraging rather than as oviposition sources. To identify potential compounds that might be useful in developing moth attractants, especially for females, collections of volatiles were made from plants that were attractive to moths in the olfactometer. Green leaf volatiles, floral volatiles, aromatic compounds, monoterpenes and sesquiterpenes were found. We propose an approach to developing synthetic attractants, here termed 'super-blending', in which compounds from all these classes, which are in common between attractive plants, might be combined in blends which do not mimic any particular attractive plant.     

Specificity of sex pheromones, the role of host plant odour in the olfactory attraction of males, and mate recognition in the aphid Cryptomyzus

Abstract. Choice tests in an olfactometer showed that males of the aphid Cryptomyzus galeopsidis spent more time in the odour of conspecific females than of a sibling species or a more distantly related Cryptomyzus species, which all co-occur on red currant. Analysis of the pheromone-emissions of females showed the presence of a nepetalactol and a nepetalactone, two known aphid sex pheromone components. However, no interspecific difference in the ratio of the components ( c. 30:1) was found, and this ratio did not change as the females aged. Males were only slightly attracted by a synthetic 30:1 mixture of these two components, and in a choice test preferreď the odour of conspecific females. The sex pheromone of Cryptomyzus spp. must thus consist of more than two components, the first time this has been reported for aphids. Host plant odour did not attract males or enhance the activity of the pheromone.     

Host location and discrimination mediated through olfactory stimuli in two species of Encyrtidae

The role of olfactory stimuli in host detection and evaluation was studied in two encyrtid Hymenoptera. The first, Epidinocarsis lopezi De Santis, is a monophagous parasitoid of the cassava mealybug Phenacoccus manihoti Matile-Ferrero, itself feeding exclusively on cassava, Manihot esculenta Crantz. The second, Leptomastix dactylopii Howard, is a monophagous parasitoid of the Citrus mealybug, Planococcus citri Risso, but this latter is highly polyphagous. The behaviour of females of both parasitoids (attaction and locomotion) was compared in a tubular olfactometer for the odours of their respective hosts on cassava and poinsettia. Tests were made using: 1) healthy host-plant alone; 2) host-plant infested with unparasitized mealybugs; 3) unparasitized mealybugs only; 4) host-plant infested with parasitized mealybugs and 5) parasitized mealybugs only. Only E. lopezi was attracted by the odour of the host-plant alone, but both species were attracted by the odour of an infested host-plant and that of unparasitized mealybugs. The odour of parasitized mealybugs, alone or on host-plant, induced an undirected activity. The attraction of E. lopezi to the odour of the host-plant alone could be linked to the monophagous diet of its host, whereas the attraction of the two species of parasitoids to the odours of infested host-plants and unparasitized mealybugs could be due to the fact that both parasitoids are specialists. The behavioural response of both species to the odour of parasitized mealybugs revealed a new aspect in host discrimination: the identification of parasitized hosts could be partly mediated through olfactory stimuli, and not only through gustatory stimuli.     

Odor learning byLeptopilina boulardi,a specialist parasitoid (Hymenoptera: Eucoilidae)

We studied odor learning in Leptopilina boulardi,a specialist larval parasitoid of Drosophila melanogaster.The behavioral responses of differently experienced females to an artificial odor (Must de Cartier, Paris) were analyzed using a fourarmed airflow olfactometer. The responses of females with an oviposition experience in the presence of the perfume were compared with those of four control groups. As controls we used naive females, females with an oviposition experience in the absence of odor, females which had been previously exposed to perfume but without an oviposition experience, and females with an oviposition experience which also had been exposed to perfume but not at the same time. The results demonstrate that a specialist such as L. boulardican learn very well to respond to an artificial odor by associating this odor with a reward, i.e., an oviposition. The four control groups responded more or less in a similar way.     

Physical and chemical plant characters inhibiting the searching behaviour of shape Trichogramma chilonis

Several plant characters are known to affect the searching behaviour and parasitization efficiency of Trichogramma spp. (Hymenoptera: Trichogrammatidae). In this study, plant characters contributing to the low Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) egg parasitism levels on pigeonpea (Cajanus cajan (L.) Millspaugh) were investigated. The efficiency of T. chilonis on pigeonpea was dependent on the plant structure on which the host eggs were found. In a cage experiment, more than 55% of eggs placed on leaves were parasitized, while 1% of eggs on calyxes and no eggs on pods were parasitized. In a filter paper bioassay, parasitoids were deterred by acetone and hexane surface extracts from pigeonpea pods but showed no response to water extract. The searching behaviour of the parasitoids was not affected by different solvent extracts from the surface of pigeonpea leaves. In a four-armed airflow olfactometer, T. chilonis was repelled by volatiles from pigeonpea pods but showed no response to volatiles derived from hexane extract of pod surfaces. Volatile infochemicals and hexane surface extracts from pods of two wild Cajanus species, C. scarabaeoides (L.) Thours and C. platycarpus (Bentham) van der Maesen, were similarly deterrent to T. chilonis. The movement of the parasitoids on pigeonpea pods and calyxes was inhibited by long trichomes and wasps were trapped by sticky trichome exudates. Parasitoids walked significantly faster on leaves than on pods. The walking speed on both pods and leaves increased significantly after washing with hexane. The results presented in this paper show that the plant growth stage and the plant structures preferred by H. armigera for oviposition are the least suitable for T. chilonis, contributing to the low parasitoid efficiency on pigeonpea.     

The major histocompatibility complex and the chemosensory signalling of individuality in humans

The chemosensory identity of mice and rats is determined partly by polymorphic genes of the major histocompatibility complex (MHC). In inbred strains of mice, as well as in seminatural populations, MHC‐associated mating preferences selectively influence reproductive success, thus serving to promote heterozygocity in the MHC. In order to determine whether MHC‐associated chemosignals are present in humans, two studies were conducted. In a first study, olfactory identification of MHC‐associated chemosignals was conducted on 12 trained rats' responses to the urine odors of humans. In a second study, MHC‐associated olfactory cues in humans were analyzed by means of gas chromatography. The results indicate that the urine odors of humans are associated with the MHC and demonstrate that the profile of volatile components in the urine odors shows some association with the MHC. Furthermore, results show that a profile of some specific components, as well as a few ubiquitous volatiles, constitutes MHC‐associated odor signals in humans. This revised version was published online in July 2006 with corrections to the Cover Date.     

Different bioassays for investigating orientation responses of the banana weevil, Cosmopolites sordidus, show additive effects of host plant volatiles and a synthetic male-produced aggregation pheromone

Abstract Three different bioassay methods to investigate the orientation behaviour of the banana weevil, Cosmopolites sordidus (Germar) (Coleoptera: Curculionidae), to host plant volatiles and a synthetic pheromone (cosmolure+) were compared. A locomotion compensator was used to separately record walking tracks in response to three odour sources. The data show that C. sordidus uses odour‐conditioned anemotaxis in its orientation to the odour sources tested. Of the two olfactometers tested, a dual port olfactometer using a continuous airflow showed stronger discrimination by C. sordidus to the different odours compared with a double pitfall olfactometer. The results of all three bioassays indicate that C. sordidus responds in an additive way to the combination of fermentation plant volatiles and the synthetic pheromone.     

Diapause status of females affects attraction of male pear psylla, Cacopsylla pyricola, to volatiles from female-infested pear shoots

A companion study showed that male pear psylla, Cacopsylla pyricola (Förster) (Homoptera: Psyllidae) were attracted to volatiles from pear shoots infested with post-diapause females. The present study compared the behavioral response of males to diapause and post-diapause females. Assays were done using a Y-tube olfactometer. We collected male and female winterform psylla from pear orchards at regular intervals between late October (early diapause) and late February (post-diapause). Female-infested shoots were not attractive to males until the February samples, coinciding with ovarian maturation and onset of mating in the field. A second set of assays was done in which we manipulated diapause status in the laboratory either by exposing psylla to a long-day photoperiod or by treating insects with an insect growth regulator, fenoxycarb. In the photoperiod experiments, both short-day and long-day males preferentially selected long-day (post-diapause) females over short-day (diapause) females. Fenoxycarb-treated males preferred fenoxycarb-treated (post-diapause) females over untreated (diapause) females; untreated males showed no preferences. Results support observations made elsewhere that male winterform pear psylla perceive and are attracted to volatile odors associated with pear shoots infested with post-diapause females.     

Olfactory response of the anthocorid predatory bug Orius sauteri to thrips-infested eggplants

The predator Orius sauteri (Poppius) (Heteroptera: Anthocoridae) is an effective arthropod natural enemy of thrips, especially Thrips palmi Karny (Thysanoptera: Thripidae), a serious pest of vegetables. First, we studied O. sauteri's response to the odour from thrips-infested eggplant [Solamum melongena L. (Solanaceae)] in a laboratory by two-choice experiments using a Y-tube olfactometer. When detached eggplant leaves were used as odour sources, O. sauteri preferred the volatiles from uninfested leaves to clean air. Concerning preferences among differently infested leaves, O. sauteri preferred the volatiles from plants infested with 10–100 thrips per leaf to uninfested leaves, but showed no significant preference for artificially damaged leaves over uninfested leaves. Similar results were obtained when complete plants were tested as the odour source. Second, release and recapture experiments in a greenhouse, a more realistic set of conditions, were conducted to confirm whether a significant preference for infested plants occurred at similar infestation levels as in the laboratory. Trends favouring infested plants were detected at densities of five and 500 thrips per plant; however, at the five thrips per plant, this trend was due to the large deviation seen in infested plants in only one replicate. In light of the low tolerable thrips density of eggplant, it is necessary to confirm whether artificial treatments with chemicals induce the emission of herbivore-induced plant volatiles that can attract O. sauteri and prolong its residence time on the leaf.     

Responses of the stem borer larval endoparasitoid Cotesia flavipes (Hymenoptera: Braconidae) to plant derived synomones: Laboratory and field cage experiments

Laboratory and field cage experiments investigated the response of females of the stem borer larval endoparasitoid Cotesia flavipes to two synthetic synomone components, the terpenoid (E)-β-farnesene and the green leaf volatile, (Z)-3-hexenyl acetate, both compounds identified previously in headspace volatiles of maize plants damaged by stem borer (Chilo partellus). In dose response tests performed in a Y-tube olfactometer, parasitoids were significantly more attracted to the arms bearing 10 or 15 µg of (Z)-3-hexenyl acetate and (E)-β-farnesene than to the control arm. (E)-β-Farnesene was as attractive as the essential oil from the plant Hemizygia petiolata (Lamiaceae) rich in the same compound (80% relative amount). The plant essential oil elicited responses from females of the parasitoid comparable to those elicited by two positive controls, stem borer larval frass and adult parasitoid diet (20% honey solution), tested in the laboratory assays. In field cage trapping experiments, captures in traps baited with the terpenoid, the plant essential oil, (Z)-3-hexenyl acetate and the control of 20% honey solution, were not significantly different relative to captures in unbaited traps. Addition of the green leaf volatile (Z)-3-hexenyl acetate to the plant essential oil to yield a 1:1 two-component blend captured significantly more female parasitoids than traps baited with either of the two components alone. The results show that blends of green leaf volatiles and sesquiterpenoids may have potential in monitoring C. flavipes populations in the field.