Pheromone-mediated Flight Research Articles

Sensory Adaptation of Antennae and Sex Pheromone-Mediated Flight Behavior in Male Oriental Fruit Moths (Leptidoptera: Tortricidae) After Prolonged Exposure to Single and Tertiary Blends of Synthetic Sex Pheromone

Sensory adaptation has been measured in the antennae of male Grapholita molesta (Busck) after 15 min of exposure to its main pheromone compound (Z)-8-dodecen-1-yl acetate (Z8-12:OAc) at the aerial concentration of 1 ng/m(3) measured in orchards treated with pheromone for mating disruption. Exposing males to this aerial concentration of Z8-12:OAc for 15 min, however, had only a small effect on their ability to orientate by flight to virgin calling females in a flight tunnel. Experiments were undertaken to determine if exposure to the main pheromone compound in combination with the two biologically active minor compounds of this species, (E)-8-dodecen-1-yl acetate (E8-12:OAc) and (Z)-8-dodecen-1-ol (Z8-12:OH) would induce greater levels of sensory adaptation and have a greater effect on male sexual behavior. The exposure of male antennae to 0.5 g/m(3) air of one of the three pheromone compounds induced sensory adaptation to this compound and to the other two pheromone compounds demonstrating cross adaptation. Average percentage sensory adaptation to a pheromone compound was similar after 15 min of exposure to 1 ng/m(3) air of Z8-12:OAc, or to 1 ng/m(3) air of a 1:1:1 or 93:6:1 blend of Z8-12:OAc, E8-12:OAc, and Z8-12:OH. The exposure of males to 1 ng/m(3) air of Z8-12:OAc or the two ratios of Z8-12:OAc, E8-12:OAc, and Z8-12:OH for 15 min had no effect on their ability to orientate to a virgin calling female. The implications of these results for the operative mechanisms of sex pheromone-mediated mating disruption of this species are discussed.

Listening in Pheromone Plumes: Disruption of Olfactory-Guided Mate Attraction in a Moth by a Bat-Like Ultrasound

Nocturnal moths often use sex pheromones to find mates and ultrasonic hearing to evade echolocating bat predators. Male moths, when confronted with both pheromones and sound, thus have to trade off reproduction and predator avoidance depending on the relative strengths of the perceived conflicting stimuli. The ultrasonic hearing of Plodia interpunctella was investigated. A threshold curve for evasive reaction to ultrasound of tethered moths was established, and the frequency of best hearing was found to be between 40 and 70 kHz. Flight tunnel experiments were performed where males orienting in a sex pheromone plume were stimulated with 50 kHz pulses of different intensities. Pheromone-stimulated males showed increased defensive response with increased intensity of the sound stimulus, and the acoustic cue had long-lasting effects on their pheromone-mediated flight, revealing a cost associated with vital evasive behaviours.

Effect of a vertical object on the sex pheromone-mediated flight and landing behaviors of the male lightbrown apple moth,Epiphyas postvittana (walker) (Lepidoptera: Tortricidae)

A small (2.5-cm-wide) vertical (10-cm-high) white object influenced the sex pheromone-mediated flight and landing behaviors of maleEpiphyas postvittana. When the vertical object was positioned on a horizontal surface to the side (3–5 cm) and upwind of a pheromone source (in the middle of the surface), the distribution of landing positions of males on the surface was different from that when the object was not present; males tended to land in positions skewed toward the side of the source that the object was on. The closer the object was positioned to the source, the greater the number of males that landed on the object (rather than on the horizontal surface). This difference in landing positions (when the object was present) corresponded with changes in the flight tracks; the tracks of males flying to the surface with an object were skewed toward the object and had higher amplitude intertrack reversal distances than the tracks of males flying to a surface without a vertical object. Positioning of a vertical object progressively upwind of the source resulted, apparently, in decreased effects on the landing (and presumably flight) behavior of males. The effect of the vertical object on the flight and landing behaviors of males corresponded largely with changes in pheromone plume structure (visualized with smoke) induced by the extra turbulence in the airflow over the source. Thus it appears that the vertical object influences the behavior of maleE. postvittana largely through the olfactory sensory modality. However, when a clear, Mylar object, in place of the white object, was placed on the surface, more males landed on the Mylar object (than did on the white object), suggesting that the vertical object may also influence the behavior of males through the visual modality.

Influence of plume structure and pheromone concentration on upwind flight of Cadra cautella males

Abstract. The effects of pheromone plume structure and its concentration on the pheromone‐mediated flight of male Cadra cautella (Lepidoptera: Phycitinae) were investigated in a laminar‐flow wind tunnel. When two C. caurella males flew simultaneously along a ribbon plume of mixed smoke and pheromone, their inflight behaviour was dependent on the instantaneous structure of the plume they encountered. When a male intercepted an intact ribbon filament, he sustained a crosswind course, whereas when he intercepted a turbulent filament (created by an upwind male fragmenting the ribbon plume), he adopted a flight course more due upwind. These results indicate that C. cautella males altered their in‐flight manoeuvres in response to instantaneous changes in the fine structure of the pheromone plume. We also demonstrated that differences in the fine structure of the plume had more influence on the flight pattern of C. cautella males than a 1000‐fold range in pheromone dose. The size of the plume was increased by adding wind deflectors upwind of the pheromone source, independent of source dosage, males following ribbon plumes flew slow zigzag tracks, whereas males following large, turbulent plumes flew directly to the source in fast, straight tracks with less counterturning.

Sex pheromone-mediated flight and landing behaviors of the European corn borer,Ostrinia nubilalis (Hübner)

The pheromone-mediated flight and landing behaviors of maleOstrinia nubilalis were studied in a wind tunnel. The pheromone source was placed in the middle of an 18 × 18-cm horizontal surface, and a smaller surface placed 4, 18, or 36 cm downwind. The smaller surface did not appear to affect significantly the flight tracks or position of landing of males on the upwind surface, and it allowed the positions and altitudes of males as they passed over the downwind surface to be estimated. The flight altitude and position of males as they passed over the downwind surface related to where males landed on the upwind surface. Regardless of the downwind position of the downwind surface, most males flew over its center (i.e., in line with the source) and landed in line with the source on the upwind surface. When a small 2.5 × 10-cm vertical object was placed on the upwind surface, just upwind and to one side of the source, males flew over the downwind surface in positions skewed toward the vertical object and in broader distributions than for the comparable situation without an object: males landed on the upwind surface on positions skewed toward, or on, the object and with a broader distribution (laterally). Flight altitude also corresponded with landing position. Thus, when there was no vertical object, most males flew just above the downwind surface and landed on the downwind edge of the upwind surface. In contrast, with the vertical object, males flew significantly higher and tended to land past the downwind edge of the upwind surface. With a taller object (20 cm), males flew even higher, past the downwind edge and most landed on the vertical object. These data show the close relationship between flight and landing behaviors of maleO. nubilalis and suggest that flight maneuvers that determine track and altitude largely govern where a male lands.

Factors influencing the landing of maleEpiphyas postvittana (Walker) exhibiting pheromone-mediated flight (Lepidoptera: Tortricidae)

The effects of changes in various visual and olfactory properties of a white card surface on the landing position of male Epiphyas postvittanaexhibiting pheromone-mediated flight were studied in a wind tunnel. Males landed predominantly at the most downwind position of a surface in line with the pheromone source, regardless of the strength of the source. The position on the surface that males landed was strongly influenced by visual factors. The landing position of males appeared to be influenced by visual cues along all three axes of the surface. Decreases in either the dimension horizontally perpendicular to the wind direction or the vertical dimension resulted in greater numbers of males landing farther upwind on the surface than the downwind edge. Visual changes in the axis along the wind direction also affected the position at which males landed. For example, when presented with two white card surfaces with a 4- cm gap between them, males tended to land on the downwind edge of the upwind surface (on which the source was located). When the gap was bridged with clear Mylar, the landing pattern was significantly different, with the greater proportion of males landing on the downwind surface. However, when Mylar was placed on the plexiglass floor of the tunnel (in addition to bridging the gap), the landing pattern on the surface was not significantly different from that on the two surfaces without the Mylar bridge. It is suggested that during the prelanding and landing phases of pheromone-mediated flight, male moths orient to visual features of the surface containing the pheromone source rather than to visual features of the source (conspecific female moth) itself.

Sublethal Effects of Cypermethrin on Chemical Communication, Courtship, and Oviposition in the Cabbage Looper (Lepidoptera: Noctuidae)

Sublethal doses of cypermethrin had multiple deleterious effects on reproductive activities in the cabbage looper, Trichoplusia ni (Hubner). Cypermethrin-treated female moths were less likely to show calling behavior than control moths. However, no differences were observed between treated and control females in the pheromone emission rates or pheromone blend ratios from forcibly extruded glands. Thus, those treated females that did call probably emitted sufficient pheromone to attract mates. Pheromone-mediated flight was reduced in males treated with cypermethrin, which would result in a reduction in successful mate location by affected males. In observations of courtship, treatment of males with cypermethrin reduced their mating success, but identical treatment of females had no effect. Results of this research are compared with previous studies of the effects of sublethal doses of chlordimeform and methoprene on reproductive events in T. ni .

Adaptation of male moth antennal neurons in a pheromone plume is associated with cessation of pheromone-mediated flight

Recordings of the firing rates of single antennal neurons when Agrotis segetum antennac were placed 70 cm downwind of a pheromone source revealed that cells sensitive to the most volatile component adapted rapidly in a plume from a high–concentration source known from previous studies to cause in-flight arrestment of progress towards the source. No adaptation was found in response to lower-concentration plumes known to promote high levels of sustained flight to the source with little premature arrestment. Adaptation was not observed in antennal neurons of a second species, Heliothis virescens, when they were placed in plumes of this species' sex pheromone blend, regardless of the concentration. In flight-tunnel tests these same pheromone sources evoked high levels of source location with little arrestment. These results indicate that adaptation or attenuation of antennal neuronal burst frequencies in response to rapidly arriving pheromone filaments in a plume may be important peripheral determinants of whether or not prolonged upwind flight and successful pheromone source location occurs.

Pheromone Components and Active Spaces: What Do Moths Smell and Where Do They Smell It?

The pheromone-mediated flight behavior of male Oriental fruit moths was observed in the field to test the hypothesis that male activation far downwind of a female is initiated by the major, or most abundant, component of the pheromone blend. Males responded at significantly greater distances to the three-component pheromone blend produced by females than to the major component alone or to either binary mixture containing the major component and one minor component. These results support the alternative hypothesis that the active space of a multicomponent pheromone is a function of male perception of the female-released blend of components, rather than of the major component alone, and that so-called minor components have a greater impact on male behavior farther downwind of a female than previously thought.

Pheromone-Mediated Flight by Male Plodia interpunctella (Hübner) (Lepidoptera: Pyralidae)

Flights and captures of male Plodia interpunctella (Hubner) were observed in a warehouse room containing pheromone traps baited with different doses and blends of ( Z,E )-9,12-tetradecadien-1-ol (ZETOH) and ( Z,E )-9,12-tetradecadien-1-ol acetate (ZETA). Flights were divided into two categories: preapproach and approach flight, the approach being differentiated from preapproach flight by the simultaneous occurrence of increased frequency and angle of turning, decreased net velocity, and net movement toward a trap. Approaches and captures were analyzed with respect to dose and blend. Alone, ZETOH failed to elicit responses significantly different from the control (zero dose), whereas ZETA elicited approaches and landings in traps. Significantly more landings and approaches occurred with blends of ZETOH:ZETA in a 2:8 ratio than with 4:6 or 6:4 blends, but the ratio of captures to approaches did not vary among blends. The frequency of approaches and captures increased with dose but the ratio of captures to approaches decreased when trap release rates exceeded 10 ng/h. Practical and theoretical implications of the results are discussed.