Effects of sex pheromone in electrostatic powder on mating behaviour by Lobesia botrana males

Abstract:  In laboratory trials, mating behaviour of males of Lobesia botrana Den. and Schiff. (Lep., Tortricidae) was investigated after treatment with an electrostatically chargeable powder, Entostat^TM. Male moths were powdered with either blank Entostat or Entostat containing sex pheromone. Significantly more Entostat adhered to L. botrana males when sex pheromone was added to the powder compared with blank Entostat. Powdering male moths with Entostat, with or without sex pheromone, caused a significant reduction in antennal response when antennae were placed 2 cm away in downwind direction, while there was little effect when antennae were placed 25–100 cm from the EAG in downwind direction. In a flight study, powdering males of L. botrana caused significant increase in time before flight initiation and reduction in proportion of males making contact with calling females compared with untreated males. In a mating experiment in Petri dishes (to minimize required search time to locate female), significantly fewer females mated successfully (based on dissection of bursa copulatrix) when males had been treated with pheromone-loaded powder. Overall, powdering males of L. botrana caused considerable suppression of mating behaviour on various levels, and these suppressing effects were increased after adding sex pheromone to Entostat.     

Reduced antennal sensitivity, behavioural response, and attraction of male codling moths, Cydia pomonella, to their pheromone (E,E)-8,10-dodecadien-1-ol following various pre-exposure regimes

The effects of pre‐exposing male codling moths, Cydia pomonella (L.) (Lepidoptera: Tortricidae), to their pheromone (E,E)‐8,10‐dodecadien‐1‐ol (codlemone), in static and moving air, under laboratory and field conditions, on subsequent antennal sensitivity, behavioural responsiveness, and attraction to codlemone were investigated. In flight tunnel experiments, the percentage of moths wing fanning and taking flight were mostly unaffected, but upwind flight to, and contact with, pheromone sources known to elicit responses of both were shown to depend on the intensity and duration of previous exposure to codlemone and recovery time between exposure and assessment. Ten to 30‐min pre‐exposures to codlemone in static air (≈ 35 µg l^?1) not only caused a 99% reduction in attraction, but also significantly reduced electroantennogram response to codlemone. Recovery of full antennal sensitivity to codlemone took more than 1 h, but recovery of attraction took over 4 h, suggesting that habituation is also partially involved in reduced behavioural responsiveness following pre‐exposure. Seventy‐five min exposures to codlemone in moving air (5–10 cm s^?1) at rates of 0.9, 4.5, and 18 µg h^?1 from Celcon fibres caused 75, 86, and 99% disruption, respectively. However, 30–34‐h exposure of caged moths to air moving through an orchard treated with 1000 Isomate‐C® dispensers ha^?1 releasing approximately 20 µg h^?1 per dispenser during tests, had no impact on moth response in flight‐tunnel assays 30 min after removal from the orchard. In this treated orchard, catches of free‐flying moths in pheromone‐baited traps were completely inhibited. If observed mechanisms such as long‐lasting antennal adaptation or habituation of the central nervous system contribute to the disruption of pheromone communication among codling moths under field conditions, it seems unlikely that they occur following exposure to the average atmospheric concentrations of codlemone. For these effects to be important, codling moths may require close contact with pheromone sources for extended periods, or repeated close encounters.     

Higher densities of distributed pheromone sources provide disruption of codling moth (Lepidoptera: Tortricidae) superior to that of lower densities of clumped sources

Field experiments quantified the effect of synthetic pheromone release-site density and distribution on 1) orientational disruption of male codling moths, Cydia pomonella (L.) (Lepidoptera: Tortricidae), to pheromone-baited traps; and 2) fruit injury. A clustering test varied pheromone release-site density from 0 to 1,000 Isomate-C Plus dispensers per ha while maintaining the total number of dispensers at 1,000. Percentage of orientational disruption of pheromone-baited traps increased significantly as a function of increasing density of release sites. Fruit injury decreased as the density of release sites increased and was lowest in plots treated with Isomate-C Plus dispensers distributed as 1,000 point sources per ha. We also manipulated point source density of 0.1-ml paraffin-wax drops containing 5% codlemone [(E,E)-8,10-dodecadien-1-ol], and thus the total amount of pheromone deployed per hectare. The percentage of disruption of traps baited with either 1.0- or 0.1-mg codlemone lures increased with increasing density of wax drops deployed. Both trapping and field observations confirmed that wax drops were attractive to male codling moths, suggesting that disruption was mediated by competitive attraction. Development of dispensers that can be mechanically applied at high densities has potential to improve the efficacy and economics of codling moth disruption at high population densities.     

Investigations of Mechanisms of Pheromone Communication Disruption of Choristoneura rosaceana (Harris) in a Wind Tunnel

Although atmospheric treatment with synthetic pheromone is used commercially to control several lepidopteran pests, little is known about how mate-finding behavior is altered by this procedure. Mechanisms of disruption of the mate-finding behavior of Choristoneura rosaceana (Lepidoptera: Tortricidae) were examined in wind tunnel bioassays. Male moths were exposed to synthetic sex pheromone or pheromone components prior to or during upwind flight to a calling virgin female. The proportion of males successfully contacting a calling female was reduced only when the synthetic pheromone treatment was presented simultaneously. A synthetic source of the attractive four-component pheromone placed upwind of a calling female was more effective than a less attractive two-component blend in achieving disorientation. Habituation of the central nervous system does not appear to be a significant factor in disruption of mate-finding behavior of male C. rosaceana, as exposure to pheromone prior to the bioassay did not alter the proportion of males that achieved subsequent upwind flight. Disruption of mate-finding behavior of C. rosaceana in a wind tunnel is probably the result of a combination of mechanisms including adaptation of antennal receptors, camouflage of the female-produced plume, and false-trail following which contributed to the additive disruption effect observed with the most attractive four-component pheromone.     

Behavioral observations of codling moth, Cydia pomonella, in orchards permeated with synthetic pheromone

Mating disruption of codling moth, Cydia pomonella, was studied in apple orchards treated with the main pheromone compound codlemone, (E,E)-8,10-dodecadienol, and a blend of codlemone and codlemone acetate, (E,E)-8,10-dodecadienyl acetate, a strong pheromone antagonist. Codlemone alone and the pheromone/antagonist-blend had a similar effect on the behavior of males emerging into air-permeated orchards. Male flights within tree canopy and upwind orientation along tree rows were strongly enhanced by both formulations, compared to untreated plots. However, the codlemone/codlemone acetate-blend increased the rate of cross-wind and downwind flights within the orchard, compared to codlemone alone. The major difference between these two formulations was that males from nearby, untreated orchards were attracted towards orchards treated with codlemone, but not towards treatments with codlemone/codlemone acetate. Additional tests were done with an equilibrium blend of codlemone and its geometric isomers. Aerial pheromone concentrations in the orchards were recorded by the field electroantennogram technique. Decreasing pheromone concentrations towards the upper part of the tree canopy, together with the stimulation of male flight activity by synthetic pheromone explains failures to control codling moth at high population densities with current dispenser formulations.     

Flight tunnel response of codling moth Cydia pomonella to blends of codlemone,codlemone antagonists and pear ester

Upwind orientation flights of codling moth males Cydia pomonella L. to a single source of sex pheromone (E,E)‐8,10‐dodecadienol (codlemone) are significantly reduced when blending it with pheromone antagonists, either with codlemone acetate, (E,E)‐8,10‐dodecadienyl acetate, or with the codlemone isomer (E,Z)‐8,10‐dodecadienol. However, once activated by a pheromone stimulus, males no longer distinguish between a pheromone source and these antagonistic blend sources. This shows that the pheromone stimulus required for the initiation of an upwind flight response differs from the stimulus for maintaining upwind flight and landing at the source. In contrast to pheromone antagonists, males discriminate between pheromone alone and a blend source of pheromone and the plant volatile pear ester, ethyl (2E,4Z)‐2,4‐decadienoate. This indicates a difference in the detection and neural integration of pheromone and plant volatile stimuli.     

Effects of exposure to pheromone and insecticide constituents of an attracticide formulation on reproductive behavior of oriental fruit moth (Lepidoptera: Tortricidae)

The effect of exposure to both the pheromone and insecticide constituents of an attracticide formulation on subsequent pheromonal response of male oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae), was tested in several wind tunnel bioassays. Male response to the attracticide formulation was significantly reduced in all behavioral categories, including source contact 1 h after sublethal exposure (both by voluntary contact in the wind tunnel and forced application in the laboratory) to the attracticide formulation containing inert ingredients, pheromone, and insecticide. Sublethal exposure to the attracticide formulation in the laboratory (forced application) 24 h before the bioassay resulted in a significantly lower proportion of males subsequently responding to attracticide droplets in the wind tunnel. However, voluntary contact of male moths with the toxic formulation in the wind tunnel had no effect on subsequent response 24 h later. Exposure of males to different constituents of the attracticide formulation demonstrated that both pheromone and insecticide exerted effects on subsequent male pheromonal response. Exposure to the formulation containing the inert ingredients plus the pheromone (no insecticide) significantly reduced male behavioral responses to an attracticide droplet in the wind tunnel 1 h but not 24 h after exposure, compared with males treated with inert ingredients alone. Response to attracticide droplets was further reduced by exposure to the entire attracticide formulation containing inert ingredients, pheromone and insecticide at both 1 and 24 h postexposure. Similarly, males exposed to inert ingredients plus pheromone were less likely to orient to female-produced plumes 1 h but not 24 h after exposure than males treated with inert ingredients alone. Response to female-produced plumes was further reduced at 1 h but not at 24 h after exposure to the entire attracticide formulation. Mating success of males was significantly reduced by exposure to the entire attracticide formulation but not to the formulation without insecticide when placed with females 1 and 24 h postexposure. These findings suggest that sublethal poisoning of males exposed to the attracticide formulation will enhance the effectiveness of this formulation under field conditions.     

Cowpea weevil flights to a point source of female sex pheromone: analyses of flight tracks at three wind speeds

Abstract.  Two-day-old male cowpea weevils, Callosobruchus maculatus, fly upwind to a point source of female sex pheromone at three wind speeds. All beetles initiating flight along the pheromone plume make contact with the pheromone source. Analysis of digitized flight tracks indicates that C. maculatus males respond similarly to moths tested at several wind speeds. Beetles' mean net upwind speeds and speeds along their track are similar ( P  > 0.05) across wind speeds, whereas airspeeds increase ( P <  0.01) with increasing wind speed. Beetles adjust their course angles to fly more directly upwind in higher wind speeds, whereas track angles are almost identical at each wind speed. The zigzag flight paths are generally narrow compared with most moth flight tracks and interturn distances are similar ( P  > 0.05) at the wind speeds employed. The frequency of these counterturns across the wind line is almost constant regardless of wind speed, and there is little variation between individuals. The upwind flight tracks are more directly upwind than those typically seen for male moths flying upwind toward sex pheromone sources. Male moths typically produce a bimodal distribution of track angles to the left and right of the windline, whereas C. maculatus males' track angles are centred about 0°. Preliminary examination of two other beetle species indicates that they fly upwind in a similar fashion.     

Pheromone pre-exposure and mating modulate codling moth (Lepidoptera: Tortricidae) response to host plant volatiles

Abstract 1 Two codling moth Cydia pomonella kairomonal attractants, ethyl (E,Z)‐2,4‐decadienoate (pear ester) and (E)‐β‐farnesene, were tested in an insecticide‐sprayed apple orchard and an orchard treated for mating disruption with synthetic pheromone (E,E)‐8,10‐dodecadienol (codlemone). Male captures with pear ester were higher in the pheromone‐treated than in the insecticide‐treated orchard, whereas captures with (E)‐β‐farnesene were not different. Subsequent wind tunnel experiments confirmed that pre‐exposure to sex pheromone codlemone increased the behavioural response of codling moth males to pear ester. This supports the idea that male attraction to the plant volatile pear ester and sex pheromone codlemone is mediated through the same sensory channels. 2 Pear ester is a bisexual codling moth attractant and even captures of female moths were significantly increased in the pheromone‐treated orchard. In the laboratory wind tunnel, pheromone pre‐exposure had no effect on female response to pear ester, but significantly more mated than unmated codling moth females flew upwind towards a pear ester source. Differences in mating status in insecticide‐treated vs. pheromone‐treated orchards may thus account for the differences in female trap captures with pear ester. 3 These findings are important with respect to monitoring of codling moth with pear ester in mating disruption orchards. They also emphasize the importance of host plant volatiles in pheromone‐mediated mating disruption, which has been neglected to date.     

Effects of adhesive powders on the mating and flight behavior of Mediterranean fruit fly (Diptera: Tephritidae)

Powders that adhere to insect cuticle can be used as carrier particles for synthetic insecticides, entomopathogens, or pheromones in insect control systems, and insects can be lured into contact with such powder mixtures by using attractants. Secondary transfer of adhesive powders to conspecifics during social interactions has been reported; however, this transfer relies on insects leaving the source of powder and continuing normal behavior when contaminated. We examined the ability of the Mediterranean fruit fly, Ceratitis capitata Wiedemann (Diptera: Tephritidae), to fly and mate after being contaminated with one of two adhesive powders: an electrostatic wax powder, Entostat, and a proprietary metallic powder, Entomag. During continuous observations for 1 h in a flight tunnel, male C. capitata made significantly more flights than females. Treating C. capitata with either powder significantly suppressed the flight activity of male C. capitata compared with untreated controls, whereas powder treatment had a negligible effect on female flight activity. Within 1 h, male C. capitata treated with Entomag recovered normal flight activity, but Entostat-treated males were not fully recovered. Virgin male C. capitata treated with either Entostat or Entomag were able to mate with virgin female C. capitata, but the onset of mating was delayed compared with control C. capitata by approximately 1 h. Even though the effect of powder uptake on behavior seemed to be temporary, scanning electron micrograph images of treated C. capitata showed that both powders were retained for > 24 h on most body parts. The adhesive powders showed potential for use as carrier particles for pesticides, entomopathogens, or pheromones in novel C. capitata control systems.     

Photoperiod cues and the modulatory action of octopamine and 5-hydroxytryptamine on locomotor and pheromone response in male gypsy moths,Lymantria dispar

Experiments were conducted to determine whether the biogenic amines octopamine (OA) and 5-hydroxytryptamine (5-HT) exert modulatory effects on pheromone responsiveness and random locomotor activity in male gypsy moths. When injected into males, OA significantly enhanced sensitivity to pheromone, while 5-HT enhanced general locomotor activity, results that were very similar to those previously shown for the cabbage looper. Maximal effect of the amines, however, was observed when injection occurred just prior to the onset of scotophase, rather than photophase, as we had originally hypothesized for this diurnally active insect. Male gypsy moths also displayed a prominent scotophase response, with sensitivity to pheromone greater in the scotphase compared with photophase, but with the level of random locomotor activity lower in scotophase than in photophase. The upwind flight behavior of males to a pheromone source in a wind tunnel, as well as the time spent at the source, were also significantly different in the two light regimes. Furthermore, when exposed to a 1 h scotophase (instead of the normal 8), or to continuous dark conditions, while males exhibited response to pheromone and locomotor activity during the same scotophase and photophase periods as observed in a 16:8 light : dark cycle, the levels of response, as well as qualitative aspects of the upwind flight behaviors in both periods were a function of the light intensity. Our combined results suggest that male gypsy moths display a bimodal rhythm of locomotor and pheromone response over the diel cycle, with light intensity and scotophase onset providing critical cues for the expression of behaviors, as well as the modulatory action of the amines. © 1992 Wiley-Liss, Inc.     

Aim‐then‐shoot anemotaxis involved in the hopping approach of potato tuberworm moth Phthorimaea operculella toward a sex pheromone source

Male moths locate conspecific females by pheromone‐induced upwind flight maintained by detecting a visual flow, termed optomotor anemotaxis. Their behavioural pattern is characterized by an upwind surge in response to a pheromone stimulus and crosswind casting after odour loss, which is considered to be reset and restarted on receipt of another pheromone pulse. However, pheromone‐stimulated males of the potato tuberworm moth Phthorimaea operculella exhibit a series of short and straight intermittent flights, or hops, when moving upwind. It is unclear whether they navigate by employing the same behavioural pattern and wind detection mechanism as that used by flying moths. To analyze odour‐modulated anemotaxis in male potato tuberworm moths, a flat wind tunnel is constructed to give regular odour stimuli to an insect regardless of its location. Moths are subjected to pheromone pulses of different frequencies to test whether they show a behavioural pattern that is reset and restarted by a pheromone pulse. Moths on the ground are also subjected to crosswind shear to examine their detection of wind direction. Path analyses reveal that males surge upwind when they receive a pheromone pulse and exhibit casting by successive hops when they lose odour. This behavioural pattern appears to be similar to that of flying moths. When the direction of the airflow is switched orthogonally, males adjust their course angle accordingly when they are on the ground. It is suggested that, instead of optomotor anemotaxis, this ‘aim‐then‐shoot’ system aids the detection of wind direction, possibly by mechanosensory means.     

Impact of (Z)-7-dodecenol and turbulence on pheromone-mediated flight manoeuvres of male Trichoplusia ni

Abstract. Turbulence and chemical noise are two factors which may influence pheromone-mediated flight manoeuvres of a moth in natural habitats. In this study, the effects of turbulence and the behavioural antagonist (Z)-7-dodecenol on flight manoeuvres of male Trichoplusia ni (Hübner) were evaluated in a wind tunnel. Male moths increase airspeed and course angles when turbulence is increased. This leads to significant increases in the length of flight tracks, but significant reductions in the time taken to reach a pheromone source. In less disturbed pheromone plumes, distributions of course angles and track angles of male T.ni show a prominent peak centred about 0° relative to the upwind direction, indicating that moths can temporarily steer directly upwind toward a pheromone source.
When (Z)-7-dodecenol is released 10 cm upwind of a pheromone source to form an overlapping plume downwind, course angles, airspeeds and ground-speeds of male T.ni are reduced significantly compared with those in uncon-taminated pheromone plumes. This results in a longer flight time to reach a pheromone source. The decrease in flight speed would decrease the rate of contact with filaments, and thereby perhaps allow the moth to detect uncon-taminated pheromone filaments independently from filaments containing the behavioural antagonist.     

Sexual differences in the flight performance of the oriental fruit moth, Cydia molesta

Cydia molesta Busck (Lepidoptera: Tortricidae), a major pest of stone fruits and an increasingly important late season pest of apple, is predominately monitored by pheromone trapping of male moths. We investigated flight performance in relation to sex, age, and mating status using computer-linked flight mills, and also examined the relationship between female flight and reproduction. The crepuscular flight pattern of the experimental moths in relation to photointensity was very similar to that reported from field studies. Female moths significantly outperformed males in all measured flight parameters including total distance flown, distance of longest single flight, and velocity. The proportion of long-flying females (categorised as those completing an unbroken flight of greater than 1 km) was three to six times greater than that of males. Female flight performance was not related to mating status, but mated males displayed significantly greater flight than unmated males. The maximal flight period of mated females commenced on the third day after eclosion, following 30% egg deposition. Male flight was not significantly related to age. The data suggest that a limited proportion of the population, in particular females, may have the capacity to make inter-orchard flights. The limitations of monitoring C. molesta populations in apple orchards solely by pheromone trapping of males are discussed.     

Temperature modulation of behavioural thresholds controlling male moth sex pheromone response specificity

ABSTRACT. The response specificity of male Oriental fruit moths, Grapholita molesta (Busck) and pink bollworm moths, Pectinophora gossypiella (Saunders), to different blends and doses of pheromone is altered dramatically by temperature. When acclimated and tested in a flight tunnel at 20^oC males of both species exhibit a high degree of specificity, with peak response levels occurring to a narrow range of blend-dose combinations close to the natural blend. When tested at 26^oC, however, males exhibit a significantly lower degree of specificity, with peak response occurring to a broader range of treatments. The change in response specificity results from shifts in behavioural threshold effects influencing plume orientation and initiation of upwind flight, as well as from arrestment of upwind flight, occurring later in the flight sequence. The observed changes in male behaviour suggest that the effect of temperature is directly on neural pathways involved in the perception of odour, and not simply the result of an increase in motor activity or a significant change in the release rate of the pheromone. The results support the threshold hypothesis for pheromone perception (Roelofs, 1978) as a general principle in the Lepidoptera, but also show that the degree of response specificity can be significantly affected by temperature.     

Evaluating dispensers loaded with codlemone and pear ester for disruption of codling moth (Lepidoptera: Tortricidae)

Polyvinyl chloride polymer (PVC) dispensers loaded with ethyl (E,Z)-2,4-decadienoate (pear ester) plus the sex pheromone, (E,E)-8,10-dodecadien-1-ol (codlemone) of codling moth, Cydia pomonella (L.), were compared with PVC dispensers and a commercial dispenser (Isomate-C Plus) loaded with codlemone. Evaluations were conducted in replicated plots (0.1-0.2 ha) in apple, Malus domestica (Borkhausen) during both generations of codling moth from 2007 to 2009. Dispensers were applied at 1,000 ha(-1). Male captures in traps baited with virgin female moths and codlemone lures were recorded. Residual analysis of field-aged dispensers over both moth generations was conducted. Dispensers exhibited linear declines in release rates of both attractants, and pear ester was released at a significantly higher rate than codlemone during both time periods. The proportion of virgin female-baited traps catching males was significantly lower with combo dispenser TRE24 (45/110, mg codlemone/mg pear ester) during the second generation in 2007 and the combo dispensers TRE144 (45/75) and TRE145 (75/45) during the first generation in 2008 compared with Isomate-C Plus. Similarly, male catches in female-baited traps in plots treated with the combo dispensers TRE144 during the first generation in 2008 and TRE23 (75/110) during the second generation, in 2007 were significantly lower than in plots treated with Isomate-C Plus. No significant differences were found for male catches in codlemone-baited traps in plots treated with Isomate-C Plus and any of the combo dispensers. However, male catches were significantly lower in plots treated with Cidetrak CM (codlemone-only dispenser) than the combo TRE144 dispenser during both generations in 2009.     

Evaluation of dyes for marking painted apple moths (Teia anartoides Walker, Lep. Lymantriidae) used in a sterile insect release program

Abstract  Two methods of marking painted apple moth ( Teia anartoides Walker, Lepidoptera: Lymantriidae), internal and external dyes, were investigated for use with the sterile insect technique, to ensure that wild and released moths could be distinguished absolutely during the eradication program. Calco Red dye was used to mark all moths internally. It was added to artificial diet and was detectable in the adults. Dye was incorporated into the larval diet at four rates. No differences were observed in mortality of the larvae, fertility of the adult female or response of male moths to virgin females in a flight tunnel. Fluorescent powder dyes were used throughout the sterile insect program to identify irradiated moths, as well as their release location and colony. However, the powdered dyes reduced the ability of male moths to detect a component of the sex pheromone in electro-antennogram tests and to respond to a calling virgin female in flight tunnel trials. Field trials suggest that moths may subsequently overcome these negative effects and respond to female moths, as moths heavily coated in dye were recaptured.     

Pheromone-triggered orientation flight of male moths can be disrupted by trifluoromethyl ketones

In a wind tunnel trifluoromethyl ketones (TFMKs) have been found to disrupt the orientation flight of male moths to pheromone sources (virgin females or synthetic pheromone). This is demonstrated by comparison of the flight parameters of the Egyptian armyworm Spodoptera littoralis and the Mediterranean corn borer Sesamia nonagrioides, which had been topically treated with TFMKs, with those calculated for untreated insects. Inhibition occurred in all types of behavior and that of the source contact has been quantified and found to be dose-dependent. The same effect has also been noticed in Mediterranean corn borer males flying to an attraction source consisting of mixtures of (Z)-11-hexadecenyl trifluoromethyl ketone (8), a closely related analogue of the major component of the pheromone, and the natural pheromone blend. The most active TFMKs are those closest in structure to the natural pheromone, along with those chemicals which easily hydrate in solution, such as the beta-thiosubstituted derivatives. Along with the previously reported reduction of catches in the field, our results suggest the possible application of these chemicals in future new pest control strategies.     

The odour makes the difference: male moths attracted by sex pheromones ignore the threat by predatory bats

How animals adjust their foraging or mate finding behaviour according to the risk of predation has received much attention recently. However, few studies have focused on the behaviour of prey when stimuli representing different adaptive behaviours originate from different sensory modalities. Here we present data on how male moths trade off odour-mediated mate attraction and auditory-mediated predator avoidance depending on the relative quality of the mate signal. Males of Agrotis segetum (Noctuidae) and Plodia interpunctella (Pyralidae), orienting towards a sex pheromone source in a flight tunnel, were exposed to ultrasound mimicking the echolocation calls of a bat, i.e. high predation risk. Males of both species accepted the predation risk when attracted to pheromone sources of high quality (female gland extract or complete synthetic blend at high dose), manifested in similar percentages reaching the pheromone source in moths with and without ultrasound exposure. In contrast, a lower proportion of ultrasound-exposed males than unexposed ones located the pheromone source when moths were flying towards pheromone sources of low quality (incomplete synthetic blend or complete synthetic blend at low dose), due to more time invested in defence manoeuvres. Furthermore, the risk taking behaviours of these moth species seem to be similar in spite of the fact that their ultrasonic hearing systems have evolved independently.     

Interactions of mechanical stimuli and sex pheromone information in antennal lobe neurons of a male moth,<Emphasis Type="Italic"> Spodoptera littoralis</Emphasis>

Male moths respond to sex pheromone sources with up-wind flight behaviour. Localization of the odour source requires not only detection of the olfactory stimulus, but also other sensory input regarding, e.g. visual and mechanical stimuli. Thus, integration of different types of sensory input is necessary. It is, however, not known where in the central nervous system the integration of information regarding different sensory modalities takes place. Using intracellular recording and staining techniques, we investigated neurons in the antennal lobe of Spodoptera littoralis, during stimulation with a mechanical stimulus and a sex pheromone. Fifteen percent of all the neurons investigated responded to the mechanical stimulus and the majority of these neurons showed altered responses if the olfactory stimulus was added. A receptor neuron responding only to the wind stimulus was found to arborise in the antennal lobe. Most projection neurons responded with an enhanced action potential frequency to the combined stimulus. In local interneurons, enhancement, depression, or no change of the responses to the wind stimulus was found when the olfactory stimulus was added. The results suggest that neurons present in the antennal lobe integrate mechanosensory and olfactory input, possibly assisting the moths to orient during up-wind flight towards an odour source.