Response of female Cydia molesta (Lepidoptera: Tortricidae) to plant derived volatiles

Peach shoot volatiles were attractive to mated female oriental fruit moth, Cydia molesta (Busck), in a dual choice arena. No preference was observed between leaf odours from the principle host plant, peach, and the secondary host plant, apple. Twenty-two compounds were identified in headspace volatiles of peach shoots using gas chromatography-mass spectrometry. Green leaf volatiles accounted for more than 50% of the total emitted volatiles. A bioassay-assisted fractionation using different sorbent polymers indicated an attractant effect of compounds with a chain length of 6-8 carbon atoms. The major compounds of this fraction were tested either singly or in combinations for behavioural response of females. Significant bioactivity was found for a three-component mixture of (Z)-3-hexen-1-yl acetate, (Z)-3-hexen-1-ol and benzaldehyde in a 4:1:1 ratio. This synthetic mixture elicited a similar attractant effect as the full natural blend from peach shoots as well as the bioactive fraction.     

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.     

Cocoon-spinning larvae of Oriental fruit moth and Indianmeal moth do not produce aggregation pheromone

1 Mature larvae of the Oriental fruit moth (OFM) Grapholita molesta (Lepidoptera: Tortricidae) and the Indianmeal moth (IMM) Plodia interpunctella (Lepidoptera: Pyralidae) leave their food source in search of suitable pupation sites in which to spin cocoons. These sites are typically well-concealed cracks and crevices within the environment. Such cocooning behaviour is also observed in larvae of the codling moth (CM) Cydia pomonella (Lepidoptera: Tortricidae), which aggregate prior to pupation in response to a pheromone blend produced by cocoon-spinning conspecific larvae.
2 In laboratory experiments, we tested whether cocoon-spinning OFM and IMM larvae produce aggregation pheromones and whether CM larvae are cross-attracted to closely-related OFM larvae.
3 Fifth-instar OFM and IMM larvae were not attracted to, or arrested by, cocoon-spinning conspecifics. Moreover, fifth-instar CM larvae were not cross-attracted to either cocoon-spinning OFM or IMM larvae.
4 Analyses of volatiles released from cocoon-spinning OFM and IMM larvae revealed that both OFM and IMM lack components that are present in the aggregation pheromone of CM larvae. This information may help explain why CM larvae are not cross-attracted to cocooning OFM or IMM larvae.     

An airflow olfactometer for measuring olfactory responses of hymenopterous parasitoids and other small insects

ABSTRACT. A new type of airflow olfactometer is described, and results given of experiments using it to measure behavioural olfactory responses of hymenopterous parasitoids. Compared with Y-tube olfactometers it shows several advantages. In its exposure chamber four separate abutting odour fields are presented so that the test insect can readily enter and re-enter them. More than one odour (or different concentrations of one odour) can be tested at the same time, thereby providing complex preference test situations. The various behavioural measures that can be assessed in the apparatus are examined and discussed.     

Thermal response in adult codling moth

Abstract.  The thermoregulation behaviour of the adult codling moth, Cydia pomonella , is investigated in the laboratory using temperature gradient experiments. Unmated males and females are tested at dawn when moths typically move to resting sites. Mated females are tested during oviposition over a complete diurnal cycle. Temperature strongly affects microhabitat selection in adult moths. Unmated males and females prefer to rest at the low-temperature ends of temperature gradients between 15 and 32 °C. Relative humidity does not influence the thermal response in unmated females, whereas males show a less distinct temperature selection under high humidity. By contrast to unmated moths, ovipositing females prove to be highly thermophilous (i.e. they deposit the highest proportions of their eggs in the zones of highest temperatures of gradients between 15 and 36 °C). This striking discrepancy in thermal response of females between their premating and oviposition period is likely to reflect an adaptation to different selection pressures from the thermal environment. Unmated moths may benefit from low temperatures by a longer lifespan and crypsis within the tree canopy, whereas the choice of warmer oviposition sites by mated females will favour a faster development of eggs.     

Widespread reliance on olfactory sex and species identification by lyssomanine and spartaeine jumping spiders

Jumping spiders (Salticidae) are renowned for their exceptional vision, but this does not preclude use of other senses. Here we provide evidence that olfactory pheromones are widespread in the Spartaeinae and Lyssomaninae, two subfamilies regarded as basal clades within the Salticidae. Pheromone use by salticids was tested in a series of experiments: males were tested with the odour of conspecific females, heterospecific females, and conspecific males, and females were tested with the odour of conspecific males. With seven of the 29 species tested, we also tested males using the draglines of conspecific females (spider absent) as the odour source. Males of all species tested were attracted to the odour of conspecific females and to the odour of the draglines of conspecific females. There was no evidence of males responding to the odour of heterospecific females or conspecific males, or of females responding to the odour of conspecific males. These findings suggest that it is primarily males that respond to olfactory sex pheromones, consistent with the apparent trend within spiders of males more actively searching for females and females placing greater emphasis on mate‐choice decisions. Compared with most salticid groups, lyssomanines and spartaeines are unusually sedentary and this lifestyle may favour olfactory mate searching. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 664–677.     

Role of host-plant stimuli in sexual behaviour of small ermine moths (Yponomeuta)

ABSTRACT. 1. Studies involving experiments with olfactometers and field experiments have shown that the release of sex-pheromone by females of Yponomeuta spp. is stimulated by the presence of host plants.
2. While in Y.evonymellus L. less females emitted pheromone in the absence of host plants, in Y.padellus L. the initiation of pheromone release was delayed.
3. Females of Y.evonymellus, Y.cagnagellus Hüb., Y.padellus and Y.plumbellus D. & S. all preferred a place with host-plant odour to one with non-host odours for sex-pheromone release.
4. Male preference for host-plant odour was demonstrated in the laboratory only for Y.cagnagellus.
5. Trap catches in the field indicated a reinforcement of female attractiveness by host-plant stimuli.     

Behavioural responses of female Neogobius melanostomus to odours of conspecifics

The behavioural responses of reproductive and non‐reproductive female round gobies Neogobius melanostomus to water conditioned by reproductive and non‐reproductive males and females were tested. The behavioural responses of reproductive female round gobies exposed to odour of reproductive males included increased time spent near the source of the odour, elevated swimming velocities and directed movement to and around the odour source when compared with their responses to control water. These results suggested that pheromones released from reproductive males may induce spawning behaviour in reproductive females. Non‐reproductive females exposed to reproductive female odour spent significantly more time near the odour source of reproductive females compared with control water. Non‐reproductive females also showed directed movement towards and around the odour source when exposed to reproductive female odour. These results suggested that round gobies use inter‐sexual and intra‐sexual pheromones and that both sex and reproductive status are important in the detection and release of these pheromones.     

A six-arm olfactometer permitting simultaneous observation of insect attraction and odour trapping

Abstract.  Behavioural assays to study insect attraction to specific odours are tedious, time consuming and often require large numbers of replications. Olfactometer and flight tunnel tests can usually only be conducted with one or two odour sources at a time. Moreover, chemical information on the odour sources has to be obtained in separate analytical studies. An olfactometer was developed in which six odours can be tested simultaneously for their relative attractiveness while during the assays, part of each test odour can be trapped for further analyses. The effectiveness of this six-arm olfactometer was tested by observing the responses of the solitary endoparasitoid Cotesia marginiventris (Cresson) to host-induced odours from young maize plants. For statistical analyses, we used log-linear models were adapted to account for overdispersion and possible positional biases. Female wasps responded extremely well in tests where they were offered a single odour source, as well as in tests with multiple choices. The responses of wasps released in groups were the same as those released individually and it was found that females did not attract or repel each other, but males preferred arms in which females had been released. Dose–response tests with varying numbers of plants or host larvae on plants revealed that the wasps responded in a dose-related manner, thus showing that the system is well suited to measure relative preference. The clear choices of the insects amongst six possibilities provided substantial statistical power. Gas chromatographic analyses of sampled air revealed clean and effective odour trapping, which largely facilitates the comparison of results from behavioural assays with the actual blends of volatiles that were emitted by the various odour sources. Advantages and disadvantages compared to other methods are discussed.     

Host odours enhance the responses of adult Rhyzopertha dominica to male-produced aggregation pheromone

The behavioural responses of adult male and female Rhyzopertha dominica (Coleoptera, Bostrichidae) to blends of host volatiles and male-produced aggregation pheromone were observed in a four-arm airflow olfactometer. The odour sources used were five pheromone-releasing males each on a single maize grain (lower maize-volatiles blend), five pheromone-releasing males on 500 g of maize (higher maize-volatiles blend) or the host volatiles emanating from 500g of maize (maize volatiles alone). Multiple-choice tests, in which volatiles from all three odour sources were presented in the exposure chamber at the same time, were used to study odour preferences of the males and females. Both sexes showed strongest attraction to the higher maize volatile blend but there were significant sex differences in response to the odour sources. Males spent significantly more time than females in the zone with only maize volatiles, and females spent significantly more time in the higher maize-volatiles zone. However, when odour sources were offered singly, females gave numerically greater responses than males to all sources although this difference was statistically significant for only the lower maize-volatiles blend. As males are more attracted than females to host odours alone it is suggested that they may be more highly adapted to seek out new hosts while females are more inclined to locate a food source by following the pheromone signals produced by males. However, both sexes responded most strongly to the odour source comprising aggregation pheromone with the higher proportion of maize volatiles.     

Preliminary experiments on the olfactory responses of Pterostichus melanarius Illiger (Coleoptera: Carabidae) to intact plants

1 In two successive years, the olfactory responses of Pterostichus melanarius Illiger (Coleoptera: Carabidae) to intact cabbage and white clover plants were studied, using a four‐arm olfactometer. 2 The first set of experiments tested the response to the odours of cabbage, white clover and the two plants grown together against potting compost; in the second and third sets of experiments, white clover and cabbage were tested, respectively. 3 The results indicate that female P. melanarius individuals are arrested by the combined odours of cabbage and white clover, spending significantly more time in the cabbage with white clover odour field than in either the cabbage, white clover or control odour fields. 4 Similar results were not found for males. 5 Cabbage plants alone did not elicit significant responses in either sex. 6 White clover alone did not elicit significant responses by females and the responses of males were not uniform in the two successive years.     

Odour quality discrimination for behavioural antagonist compounds in three tortricid species

The antennal and behavioural response of three tortricid species (Lepidoptera: Tortricidae) to their corresponding sex pheromones and known or putative behavioural antagonists was tested by electroantennography and in field trials. The species and their pheromones and known or proposed behavioural antagonist were lightbrown apple moth, Epiphyas postvittana (Walker) [pheromone: 95% (E)‐11‐tetradecenyl acetate (E11‐14Ac) and 5% (E,E)‐9,11‐tetradecadienyl acetate (E9E11‐14Ac); antagonist: (Z)‐11‐tetradecenyl acetate (Z11‐14Ac)], codling moth, Cydia pomonella (L.) [pheromone: (E,E)‐8,10‐dodecadien‐1‐ol (codlemone); antagonist: (E,E)‐8,10‐dodecadienyl acetate (codlemone acetate)], and gorse pod moth, Cydia ulicetana (Haworth) [pheromone: (E,E)‐8,10‐dodecadienyl acetate (codlemone acetate); putative antagonist: (E,E)‐8,10‐dodecadien‐1‐ol (codlemone)]. In all three species, the antennal response to the antagonists was not significantly different from the antennal response to con‐specific sex pheromone compounds. In the field trapping experiments, significantly fewer males of all three species were attracted to the respective pheromone when blended with the behavioural antagonist compound. However, this response varied between the species, with lightbrown apple moth and codling moth showing stronger responses to the antagonist compounds than gorse pod moth. Both lightbrown apple moth and codling moth males were able to discriminate between pure pheromone and pheromone blended with the antagonist when placed in traps side‐by‐side separated by ca. 10 cm. The presence of the behavioural antagonist not only affected the catch of males of both species within their own traps but also affected the catch in the neighbouring trap that contained con‐specific sex pheromone; the catch of gorse pod moth was not reduced by the presence of codlemone in the neighbouring trap. These results suggest that strong behavioural antagonists such as codlemone acetate for codling moth and Z11‐14Ac for lightbrown apple moth induce their inhibition effect at a substantial distance downwind from the odour source; however, most of those males that were able to overcome this inhibition effect at the early stage of orientation to odour source, were able to discriminate between the pheromone source and the pheromone source admixed with behavioural antagonist. Moderate behavioural antagonists such as codlemone for gorse pod moth did not elicit a discrimination effect.     

Olfactory cues in discrimination among individuals in dominance hierarchies in the cockroach, Nauphoeta cinerea

ABSTRACT. The purpose of this study was to explore the olfactory recognition mechanisms used by individual subordinate male cockroaches, Nauphoeta cinerea (Olivier), in a dominance hierarchy. Results of two independent sets of experiments, one using an olfactometer and the other using an arena, suggest that olfactory cues are important to males. Subordinate males did not avoid dominant male odour in an olfactometer but could discriminate between the odours of familiar and unfamiliar individual dominant males, preferring the odour of the dominant male with which they had previous social contact. Since test subordinates did not avoid dominant male odours in the olfactometer, it is possible that the distance an individual remains from a dominant is important. To test this, individual dominant and subordinate males were tethered in the centre of an arena. Subordinate males were introduced into the arena and allowed to approach the tethered male. They approached tethered dominants and subordinates differently, and tended to keep away from the dominants. Tethered males unfamiliar to the test animals were avoided as well.     

Meadow Voles (Microtus pennsylvanicus, Arvicolidae) Over-mark and Adjacent-mark the Scent Marks of Same-sex Conspecifics

Scent over-marking occurs when an animal deposits its scent mark on top of the scent mark of a conspecific; adjacent-marking occurs when an animal deposits its scent mark next to the scent mark of a conspecific. Given that male rodents usually scent mark more than females and that animals spend more time investigating the odor of the top-scent donor of an over-mark, I tested the following three hypotheses. First, male meadow voles deposit more scent marks than female meadow voles. Second, male meadow voles will deposit more over-marks and adjacent-marks in response to the scent marks of a same-sex conspecific than females would. Third, meadow voles spend more time investigating the odor of the second vole placed in the arena than that of the first vole placed in the arena. To test these hypotheses, two age-matched, like-sex conspecifics (first vole and second vole) were placed successively into an arena in which they were allowed to freely explore and scent mark for 15 min. The first hypothesis was not supported. The first and second vole, independently of sex, deposited a similar number of scent marks. The second hypothesis was also not supported by the data: more conspecific scent marks were over-marked by the second female than by the second male. The third hypothesis was supported by the data. After investigating a scented arena, males and females spent more time investigating the odor of the second vole than that of the first vole. Sex differences in scent-marking behaviors of meadow voles are unlike those reported for other species of rodents.     

Apple and peach fruit volatiles and the apple constituent butyl hexanoate attract female oriental fruit moth, Cydia molesta, in the laboratory

Volatiles emitted from immature and mature peach and apple fruits were all attractive to mated female oriental fruit moth, Cydia molesta (Busck), in a dual choice arena. Females did not discriminate between odours emitted by these two major host plants. The same natural blends were behaviourally ineffective for virgin females. A major component of apple fruit volatiles, butyl hexanoate, also attracted female C. molesta. Mated females were attracted to two medium dosages, while virgin females responded positively to the lowest of the five dosages tested. The time course of the captures of the moths shows a diurnal activity cycle known from the field. The possible implications of a semiochemical which attracts females are discussed in the context of previous findings that gravid females may immigrate from peaches into apple orchards particularly in the later phase of the season.     

Love is in the air: olfaction-based mate-odour identification by jumping spiders from the genus Cyrba

Jumping spiders (Salticidae) are known for having good eyesight, but the extent to which they rely on olfaction is poorly understood. Here we demonstrate for the first time that olfactory pheromones are used by two species from the salticid genus Cyrba (C. algerina and C. ocellata). Using a Y-shape olfactometer, we investigated the ability of adult males and females of both species to discriminate between mate and non-mate odour. A hidden spider or a spider’s draglines (no spider present) were used as odour sources. There was no evident response by females of either Cyrba species to any tested odour. Males of both species chose odour from conspecific females, or their draglines, significantly more often than the no-odour control, but there was no evident response by males to any of the other odours (conspecific male and heterospecific female). Our findings demonstrate that C. algerina and C. ocellata males can make sex- and species-specific discriminations even when restricted to using olfaction alone. Also, by showing that draglines can be a source of olfactory pheromones, our findings illustrate the difficulty of ruling out olfaction when attempting to test for chemotactile cues.     

Pea plant volatiles guide host location behaviour in the pea moth

Identification of plant volatiles that attract mated insect females for oviposition can provide important information about plant–insect relationships that can be used to develop pest control strategies involving manipulation of the female host search. Our study represents a first step towards identifying volatiles that affect the host location behaviour of the pea moth Cydia nigricana. The behaviours of virgin and mated males and females were analysed in cage experiments testing a two-choice situation at close range and in wind tunnel experiments evaluating upwind orientation over a distance. In both experimental setups, flowering pea plants constituted the most attractive phenological stage for mated females, with 58 % landing on such plants in the wind tunnel. Testing headspace extracts of different phenological stages of pea and of detached pea buds and flowers in the wind tunnel, mated females showed the highest landing responses to volatiles during flower development (budding 42 % and flowering 56 %) and from detached buds (46 %) and flowers (66 %). Volatile compounds collected from the various phenological stages of pea were analysed by gas chromatography–mass spectrometry, and the antennal responses to these headspace collections were evaluated by gas chromatography–electroantennography. Ten antennally active compounds were identified, nine of which were present in the headspace extracts of the whole pea plants at all tested phenological stages and in detached buds and flowers. Overall, our results demonstrate a clear link between host plant phenology, the corresponding plant odour, and the behaviour of mated C. nigricana females.     

Evidence of a rheotactic component in the odour search behaviour of freshwater eels

The detection of food odour by the freshwater eels, Anguilla australis and Anguilla dieffenbachii released a behavioural response to flow that resulted in direct upstream movement toward the odour source. Changes in various orientation parameters were observed as eels neared the source. Eels had substantially lower swimming velocities and considerably more variable heading and course angles close to the odour source (≤0·9 m) than further away (>0·9 m). Observed changes in orientation parameters were primarily due to changes in the behaviour of a searcher following odour loss. Cross-stream movements were initiated when the eel moved beyond the lateral margins of the odour plume. The behavioural switch from odour-conditioned rheotaxis to cross-stream casting following odour loss occurred more frequently close to the odour source where the plume was most narrow. Odour-conditioned rheotaxis enables the searcher to move quickly and efficiently toward the odour source without the need to extract directional information from a highly intermittent and complex chemical signal.     

Close encounters: contributions of carbon dioxide and human skin odour to finding and landing on a host in Aedes aegypti

In a wind‐tunnel study, the upwind flight and source location of female Aedes aegypti to plumes of carbon dioxide (CO₂) gas and odour from human feet is tested. Both odour sources are presented singly and in combination. Flight upwind along the plumes is evident for both CO₂ and odour from human feet when the odours are presented alone. Similarly, both odour sources are located by more than 70% of mosquitoes in less than 3 min. When both CO₂ and odour from human feet are presented simultaneously in two different choice tests (with plumes superimposed or with plumes separated), there is no evidence that females orientate along the plume of CO₂ and only a few mosquitoes locate its source. Rather, the foot odour plume is navigated and the source of foot odour is located by over 80% of female Ae. aegypti. When a female is presented a plume of CO₂ within a broad plume of human foot odour of relatively low concentration, the source of CO₂ is not located; instead, flight is upwind in the diffuse plume of foot odour. Although upwind flight by Ae. aegypti at long range is presumably induced by CO₂ and the threshold of response to skin odours is lowered, our findings suggest that, once females have arrived near a prospective human host, upwind orientation and landing are largely governed by the suite of human odours, whereas orientation is no longer influenced by CO₂.     

Visual and olfactory cue interaction in resource-location by the blowfly, Lucilia sericata

Abstract. The interactions between olfactory and visual cues in the landing responses of the blowfly Lucilia sericata (Meigen) (Diptera: Calliphoridae) were examined in a laboratory wind tunnel. The odour of liver and sodium sulphide solution released at 1 L/min, resulted in a greater number of landings, closer to, but downwind of, the central odour release point, than when odour was absent. Three-day-old protein-deprived females landed in greater numbers than protein-fed female flies of the same age; no difference existed between 6-day-old protein-fed or protein-deprived females. Six-day-old, protein-fed flies landed closer to the odour source than did 3-day-old, protein-fed flies. In the presence of odour, flies landed closer to the central release point when it was accompanied by a visual cue. No such effect of the visual cue was evident in the absence of odour. When a plain white sticky-surface (25 × 25 cm) was paired with an identical white surface plus odour, approximately equal numbers of flies landed when the surfaces were placed together; increasingly higher numbers of flies landed on the surface with the odour cue when the distance between the surfaces was increased. When a white surface with a visual cue was presented with the plain white surface plus odour, more flies landed on the white surface with the visual cue than on the plain white surface with odour when they were close together. However, as these two surfaces were moved apart, flies landed increasingly more on the surface with the odour than the surface with the visual cue. The results show that while odour cues may enhance the induction of landing by female L. sericata , visual cues are important when selecting a final landing site.