Activation, orientation and landing of female Culex quinquefasciatus in response to carbon dioxide and odour from human feet: 3-D flight analysis in a wind tunnel.

This study investigated the interaction between carbon dioxide (CO(2) ) and human foot odour on activation, upwind orientation and landing of host-seeking female Culex quinquefasciatus (Say) (Diptera: Culicidae) in a wind tunnel. More mosquitoes landed on warmed glass beads coated with foot odour than on clean beads; adding a plume of 4% CO(2) did not influence the proportion of mosquitoes landing. A second experiment used 3-dimensional video tracking to assess flight performance. Activation was more rapid with CO(2) and with CO(2) + foot odour than with clean air or with foot odour alone. Upwind flights were fastest with CO(2) and with clean air, and slowest with foot odour; the CO(2) + foot odour treatment overlapped the previous three treatments in significance. Flight headings tended more towards due upwind with CO(2) and with clean air than with CO(2) + foot odour or with foot odour alone. In both experiments, many mosquitoes flew upwind in clean air. There was little evidence of females changing course upon entering or exiting the CO(2) plume or reacting to foot odour during flight.

Efficiency of CO2 -baited CDC miniature light traps under semi-field conditions and characterizing response behaviors of female Aedes aegypti (Diptera: Culicidae).

Aedes aegypti (L.) (Diptera: Culicidae) is an important vector of viruses causing dengue, Zika, chikungunya, and yellow fever and as such is a threat to public health worldwide. Effective trapping methods are essential for surveillance of both mosquito species and disease presence. The Centers for Disease Control Miniature Light Trap (CDC-MLT) is an updated version of the New Jersey light trap, which was developed early in the 20th century. This trap is widely reported as being less successful for Ae. aegypti than for other mosquito species, although the reason for this is unclear. This trap has engendered more Ae. aegypti-tailored designs that still represent the basic design model. The efficiency of the CDC-MLT alone and with CO2 was tested under semi-field conditions and the behavior of responding female Ae. aegypti was characterized. The CDC-MLT alone failed to capture any mosquitoes and with CO2 the capture efficiency was less than 2%. Understanding the behaviors that mosquitoes exhibit while encountering a particular trap design or trapping concept may suggest trap improvements to increase capture efficiency. Moreover, this work contributes to our understanding of mosquito host-seeking behavior.

Hydrocarbon sex pheromone in tiger moths (Arctiidae).

2-Methylheptadecane is a sex pheromone compound in many sibling species of the Holomelina aurantiaca complex, in Holomelina laeta, and in Pyrrharctia isabella, which are all arctiids. Habitat preference, temporal distribution , and differing diurnal cycles help effect reproductive isolation among the species, but secondary sex pheromone chemicals are also suggested by the field studies.

Reinvestigation of oak leaf roller sex pheromone components and the hypothesis that they vary with diet.

The sex pheromone of the oak leaf roller, Archips semiferanus, was identified as a specific blend (67:33) of trans-11- and cis-11-tetradecenyl acetates. The pheromone blend of females from a semisynthetic diet and from three oak species did not vary significantly. Males from these diets responsded in the laboratory and the field only to treatments approximating the 67 : 33 blend. These findings conflict with the hypothesis that the composition and perception of month sex pheromones vary with slight changes in diet.

European corn borer: pheromone polymorphism or sibling species?

Electrophoretic analyses of the (Z) and (E) pheromone-attracted males of Ostrinia nubilalis (Hübner), the European corn borer, in an area of coexistence indicate that these strains are not freely interbreeding. Although the populations are morphologically indistinguishable, studies of allozyme, pheromone, and hybridization suggest that the (Z) and (E) entities are genetically differentiated, perhaps to the status of semi- or sibling species.

Genetic independence of female signal form and male receiver design in the almond moth, Cadra cautella.

Efficient signalling requires coordination of signal form and receiver design. To maintain signal function, parallel changes in signaller and receiver traits are required. Genetic correlation and co-evolution among signal and response traits have been proposed to preserve signal function (i.e. coordination) during the evolution of mate recognition systems. Empirical studies have provided support for both mechanisms; however, there is debate regarding the interpretation of some of these studies. Tests for a genetic correlation typically hybridize divergent signalling systems and look at hybrid signal form and receiver design, or impose artificial selection on signal form and look for an indirect response to selection in receiver design. Some of the hybridization studies did not achieve reassortment of genes from the parental types, whereas some of the artificial selection studies incorporated random mating in their designs. As a result of these limitations, the hybridization studies cannot discriminate between genetic correlation and co-evolution with primarily additive genetic effects underlying signal and response traits. Similarly, the artificial selection experiments cannot discriminate between genetic correlation because of linkage disequilibrium and co-evolution. This study examined the mating preferences of male almond moths, Cadra cautella, before and after female moths were artificially selected (using a design incorporating assortative mating) for novel pheromone blend ratios. Our results demonstrate the absence of a genetic correlation between signal and response traits in the almond moth.

Utilization of pheromones in the population management of moth pests.

Pheromones are substances emitted by one individual of a species and eliciting a specific response in a second individual of the same species. In moths (Lepidoptera) generally females lure males for mating by emission of a sex attractant pheromone comprised of either one or more components. Since 1966 the identification of the pheromone blends of many moth pests has allowed investigations into the use of these messengers for population manipulation. Pheromone-baited traps may be used both to detect pest presence and to estimate population density, so that conventional control tactics can be employed only as required and timed precisely for maximum effectiveness. Attractant traps also can be utilized for direct population suppression when the traps are deployed at a density effective in reducing mating success sufficiently to achieve control. A third use pattern of pheromones and related compounds is disruption of pheromone communication via atmospheric permeation with synthetic disruptants. The behavioral modifications involved in disruption of communication may include habituation of the normal response sequence (alteration of the pheromone response threshold) and "confusion" (inability of the organism to perceive and orient to the naturally emitted lure). Disruption of communication employing the natural pheromone components as the disruptant has been most successful, although nonattractant behavioral modifiers structurally similar to the pheromone components also may prove useful. Possible future resistance to direct pheromone manipulation may be expected to involve the evolution of behavioral and sensory changes that minimize the informational overlap between the natural pheromone system and the pheromone control technique.

Orientation of Culex mosquitoes to carbon dioxide-baited traps: flight manoeuvres and trapping efficiency.

Females of Culex quinquefasciatus Say and Culex tarsalis Coquillet (Diptera: Culicidae) in the host-seeking stage were released and video recorded in three dimensions in a large field wind tunnel as they flew to four kinds of CO2-baited mosquito traps. The trapping efficiency (number of mosquitoes approaching compared to the number caught) was determined for each trap type. The Encephalitis Virus Surveillance (EVS), Mosquito Magnet Freedom (MMF) and Mosquito Magnet Liberty (MML) traps captured only 13-16% of approaching Cx. quinquefasciatus females, whereas the Mosquito Magnet-X (MMX) trap captured 58%. Similar results were obtained for Cx. tarsalis. Orientation behaviour and flight parameters of mosquitoes approaching the four traps were compared. Mosquitoes spent the most time orienting to the EVS trap. Flight speed decreased as mosquitoes entered the vicinity of each trap and a large portion of their time was spent within 30 cm downwind of the traps. Flights became highly tortuous downwind of the poorly performing traps and just upwind of the MMX trap. Differences between traps and possible explanations for the superior performance of the MMX trap are considered.

Odour plumes and odour-mediated flight in insects.

Flying insects often follow odour plumes to find resources. Some insects may employ an 'aim-and-shoot' strategy using mechanoreceptors before flight to determine wind direction. Once airborne, insects must use optomotor anemotaxis to set a course upwind. This mechanism uses a visual appraisal of how wind modifies the insect's path. A straight upwind course yields a front-to-rear image flow directly below the insect. Details of this process in male moths flying to female pheromone have emerged mainly from wind tunnel studies. Loss of the pheromone triggers 'casting', or wide lateral excursions without upwind progress, whereas contact with a plume usually induces a zigzag path upwind. The temporally regular counterturns in casting and zigzagging seem to be generated by a central programme. Brief contact with a filament of odour induces a heading towards upwind, and an optimal rate of encounter promotes a rapid, straight upwind course. Other insects, such as parasitoid wasps seeking a host and tsetse flies seeking a blood meal, seem not to have a temporally regular pattern of counter-turns and often fly straight upwind. The availability of visual cues from the odour source itself, the aerial distribution of odour set by turbulent diffusion, and light and wind levels all influence the success of these manoeuvres.

Rate of release of spruce budworm pheromone from virgin females and synthetic lures.

The rate of pheromone [(E)- and (Z)-11-tetradecenal] release from calling virginChoristoneura fumiferana females and synthetic lures was determined in both static and aerated atmospheres. In a static system ca. 2 ng/hr was recovered per female. Owing to the > 75% adsorption onto the females' bodies in static atmospheres, the actual release rate has to be corrected to roughly 9-27 ng/hr, depending on the percentage adsorbed. In the air-flow system, females were found to release between 4 and 20 ng/hr. On a 16∶8 light-dark cycle, calling began 1-2 hr before lights-off and continued nonstop until lights-on. Pheromone was emitted throughout calling, while no pheromone was detected during the noncalling periods.

Role of (Z)- and (E)-11-tetradecenyl acetate pheromone components in the sexual behavior of the (Z) strain of the european corn borer,Ostrinia nubilalis (Lepidoptera: Pyralidae).

A glass tube olfactometer bioassay was used to examine pheromone response of males of the (Z)-pheromone strain ofOstrinia nubilalis (Hubner). The presence of (E)-11-tetradecenyl acetate at the natural ratio to (Z)-11-tetradecenyl acetate (97∶3; Z∶E) did not consistently elevate wing-fanning, upwind walking, or clasper extrusion over (Z)-11-tetradecenyl acetate alone. This bioassay did not reveal the behavioral role of (E)-11-tetradecenyl acetate.

Effect of pheromone concentration on organization of preflight behaviors of the male gypsy moth,Lymantria dispar(L.).

Male gypsy moths (Lymantria dispar L.) in a wind tunnel at 24° respond to decreasing dosages (1 µg to 0.1 pg) of (+)-cis-7,8-epoxy-2-methyloctadecane with a decrease in probability of wing-fanning, an increase in wing-fanning latency, and an increase in the number of behaviors (body jerks, antennal twitches, steps, and wing tremors) preceding fanning. Males initiating any behavior prior to wing-fanning had a 70% probability of wing-fanning and 97% of the males that wing-fanned eventually flew. The sequence of behaviors from quiescence to flight is not organized in a hierarchy, as this concept is used in ethology, nor is it dependent upon the concentration of pheromone. The time-average threshold concentration of pheromone for response of ca. 90% or more quiescent males is ca. 1.9 × 10(-17) g/cm(3) over < 0.3 min.

Mate location strategies of gypsy moths in dense populations.

In high-density populations, the male gypsy moth (Lymantria dispar) "appetitive" flight is primarily vertical and within 1-50 cm of tree boles. Preceding location of a female, males land on trees or occasionally tree foliage and walk while wing-fanning. In high-density populations females may mate before calling, often prior to wing expansion or hardening. Additional matings may occur before any or full deposition of egg masses. Virgin females are not coy, whether their wings are unexpanded, or hardened and held rooflike, or whether they are calling or not. They generally mate with the first willing male and do not exercise sexual selection by an acceptance or rejection reaction. The mating structure in dense populations would seem to reduce selective pressure for female emission of and direct male anemotactic flight to attractant pheromone. A low proportion of males "search" appropriate objects, especially tree trunks, in the walking-wing-fanning state. Such males often are successful in locating virgin females before calling commences. This male strategy presumably would not be successful in low population densities.

Wing fanning as a measure of pheromone response in the male pink bollworm,Pectinophora gossypiella (Lepidoptera: Gelechiidae).

A still-air, wing fanning bioassay measured male pheromone response in the pink bollworm moth,Pectinophora gossypiella (Saunders). As pheromone concentration was increased, the duration of wing fanning increased while its latency decreased. Response duration was longest for the natural pheromone blend [44∶:56 (Z,E)- and (Z,Z)-7,11-hexadecadienyl acetates], although responses to 38% and 50%Z,E blends were not significantly briefer. Preexposure to pheromone had no effect on wing fanning in males retested after 24 hr. Wing fanning was highly correlated with upwind flight and other responses observed in a wind tunnel. Wing fanning duration provides a continuous measure of pheromone response and allows a large number of males to be tested in a short time; these two characteristics of the wingfanning assay are helpful for genetic analyses of male response.

Heritable variation in pheromone response of the pink bollworm,Pectinophora gossypiella (Lepidoptera: Gelechiidae).

Heritability of variation in male pheromone response by pink bollworm moths,Pectinophora gossypiella (Saunders), was examined using a still-air, wing-fanning bioassay. Heritability (±SE) of overall responsiveness, as measured by the mean duration of wing fanning to the blend of pheromone components produced by females [44∶56 ratio of (Z, E)- to (Z, Z)-7,11-hexadecadienyl acetate], was 0.385 ± 0.095. Heritabilities of wingfanning duration to blends with 25 and 65%Z, E isomer were 0.377 ± 0.113 and -0.145 ± 0.103, respectively. These findings indicate an asymmetry in the genetic component of variation in response to pheromone blends with high and low proportions of theZ, E isomer. An index of response specificity for individual males was developed based on the response to an off-blend (either 25 or 65%Z, E isomer) relative to the response to the 44%Z, E blend. Heritabilities of response specificity were 0.117 ± 0.059 and -0.043 ± 0.067 for the 25 and 65%Z, E blends, respectively.

Selection for increased pheromone response in the male pink bollworm, Pectinophora gossypiella (Lepidoptera: Gelechiidae).

The genetic basis of the duration and incidence of male wing fanning to pheromone in the pink bollworm moth, Pectinophora gossypiella (Saunders), was examined by artificial selection. Using a still-air bioassay, males from a laboratory colony were selected for increased duration of wing fanning when exposed to a 65:35 blend of (Z,E)- and (Z,Z)-7,11-hexadecadienyl acetates. The mean (+/- SE) duration of wing fanning in the selected line increased from 5.4 +/- 1.4 to 17.4 +/- 2.7 s after six generations. The increase in wing fanning duration was the result of an increase in response duration among responders and not the percentage of males that responded. Realized heritability of wing-fanning duration was 0.16 +/- 0.02. The amount and ratio of pheromone produced by females in the male-selected line did not change.

Comparing the effectiveness of sexual communication disruption in the oriental fruit moth (Grapholitha molesta) using different combinations and dosages of its pheromone blend.

The relative efficacy of disruptant blends comprised of different combinations of the Oriental fruit moth's pheromone components was determined in field tests. Disruption was evaluated by comparing male moth catch at synthetic and female-baited traps in disruptant and non-treatment areas. Three atmospheric dosages of a 8-dodecenyl acetate (93.5%Z∶6.5%E) blend, representing two successive 10-fold decreases in concentration (2.5 × 10(-2) g/hectare/day to 2.5 × 10(-4) g/hectare/day) were tested alone and in combination with an additional percentage of (Z)-8-dodecen-1-ol. Male moth orientation to traps was eliminated in plots exposed to the two highest binary acetate dosages. However, significantly more males were captured in synthetic-baited traps in the lowest acetate-alone treatment, indicating a diminution of disruption efficiency. In contrast, inclusion of (Z)-8-dodecen-1-ol in the disruptant blend effected essentially complete disruption of orientation at all concentrations tested. Mating success ofG. molesta pairs confined in small cages apparently was not affected by the presence of relatively high concentrations of the binary acetate and the acetate-alcohol blends. This suggests that habituation and/or adaptation of male response, at least for comparatively "close-range" behaviors, did not occur.