Fly Upwind Research Articles

Going against the flow: bumblebees prefer to fly upwind and display more variable kinematics when flying downwind.

Foraging insects fly over long distances through complex aerial environments, and many can maintain constant ground speeds in wind, allowing them to gauge flight distance. Although insects encounter winds from all directions in the wild, most lab-based studies have employed still air or headwinds (i.e. upwind flight); additionally, insects are typically compelled to fly in a single, fixed environment, so we know little about their preferences for different flight conditions. We used automated video collection and analysis methods and a two-choice flight tunnel paradigm to examine thousands of foraging flights performed by hundreds of bumblebees flying upwind and downwind. In contrast to the preference for flying with a tailwind (i.e. downwind) displayed by migrating insects, we found that bees prefer to fly upwind. Bees maintained constant ground speeds when flying upwind or downwind in flow velocities from 0 to 2 m s-1 by adjusting their body angle, pitching down to raise their air speed above flow velocity when flying upwind, and pitching up to slow down to negative air speeds (flying backwards relative to the flow) when flying downwind. Bees flying downwind displayed higher variability in body angle, air speed and ground speed. Taken together, bees' preference for upwind flight and their increased kinematic variability when flying downwind suggest that tailwinds may impose a significant, underexplored flight challenge to bees. Our study demonstrates the types of questions that can be addressed with newer approaches to biomechanics research; by allowing bees to choose the conditions they prefer to traverse and automating filming and analysis to examine massive amounts of data, we were able to identify significant patterns emerging from variable locomotory behaviors, and gain valuable insight into the biomechanics of flight in natural environments.

Building a Uniport Olfactometer to Assess Mosquito Responses to Odors.

The uniport olfactometer behavioral assay is currently one of the most reliable single-choice methods to use to study mosquito attraction to olfactory stimuli. It allows for the reproducible calculation of mosquito attraction rate to human hosts or to other olfactory stimuli. Here, we present the design of our modified uniport olfactometer. Consistent carbon-filtered air flows through the assay, creating positive pressure that reduces odor contamination from the room. It includes a precision-milled white acrylic base to facilitate easy setup and consistent placement of the component parts. Our design can be made by a commercial acrylic fabricator or an academic machine shop. This olfactometer is designed to assess the responses of mosquitoes but could be applied to other insects that fly upwind toward an odor stimulus. In a companion protocol, we detail how to perform the experiments with mosquitoes by using the uniport olfactometer.

Quantifying Aedes aegypti Host Odor Preference Using a Two-Port Olfactometer.

Blood-feeding mosquitoes are a leading threat to global public health-vectoring dangerous infections including Zika, dengue, and malaria. Mosquitoes identify and target hosts for blood meals by using visual, thermal, and chemical cues. Here we describe an assay for measuring odor-based host-preference behavior-that is, the preferential approach toward one host over another based on differences in the volatile compounds they emit. The assay can be adapted for use with diverse odor sources, from live animals and their breath to odor-scented sleeves with controlled amounts of CO2 Mosquitoes in this assay fly upwind to within 30 cm of the odor source and then enter a small trap. We therefore believe this assay best replicates medium- to short-range host-seeking, when females approach and are preparing to land on a host animal. We also find that relative response in a two-choice test shows less trial-to-trial variation than the absolute number of responsive mosquitoes, which appears more sensitive to exogenous factors such as rearing conditions. This assay has been used to better understand mosquito host-seeking decisions, which can provide fundamental insight into the brain and behavior as well as information useful for the design of novel vector control strategies.

L-lactic and 2-ketoglutaric Acids, Odors from Human Skin, Govern Attraction and Landing in Host-Seeking Female Aedes aegypti Mosquitoes

Aedes aegypti, presented with a source of L-lactic and 2-ketoglutaric acid in a wind-tunnel bioassay, takeoff, fly upwind, and land on the odorant substrate at rates comparable to those exhibited by mosquitoes presented with a skin-odor stimulus. Addition of carbon dioxide decreased takeoff latency but was not required to elicit upwind flight nor landings. Ketoglutaric acid, a recently identified component of human skin odor, combined with lactic acid elicits the full repertoire of mosquito host-seeking behaviors.

To what extent can the tiny parasitoid wasps, Eretmocerus mundus, fly upwind?

AbstractBody miniaturization in insects is predicted to result in decreased flight speed and therefore limited ability of these insects to fly upwind. Therefore, tiny insects are often regarded as relying on passive dispersal by winds. We tested this assumption in a wind tunnel by measuring the burst speed of Eretmocerus mundus (Mercet), a beneficial parasitoid wasp with body length <1 mm. Insects were filmed flying upwind towards a UV light source in a range of wind speed 0–0.5 m/s. The Insects flew towards the UV light in the absence and presence of wind but increased their flight speed in the presence of wind. They also changed flight direction to be directly upwind and maintained this body orientation even while drifted backwards relative to the ground by stronger winds. Field measurements showed that the average flight speed observed in the wind tunnel (0.3 m/s) is sufficient to allow flying between plants even when the wind speed above the vegetation was 3–5 folds higher. A simulation of the ability of the insects to control their flight trajectory towards a visual target (sticky traps) in winds show that the insects can manipulate their progress relative to the ground even when the wind speed exceeds their flight speed. The main factors determining the ability of the insects to reach the trap were trap diameter and the difference between insect flight speed and wind speed. The simulation also predicts the direction of arrival to the sticky target showing that many of the insects reach the target from the leeward side (i.e. by flight upwind). In light of these results, the notion that miniature insects passively disperse by winds is misleading because it disregards the ability of the insects to control their drift relative to the ground in winds that are faster than their flight speed.

Semiochemicals for Thrips and Their Use in Pest Management.

Thrips (Thysanoptera) are small insects that can cause huge problems in agriculture, horticulture, and forestry through feeding and the transmission of plant viruses. They produce a rich chemical diversity of pheromones and allomones and also respond to a broad range of semiochemicals from plants. These semiochemicals offer many opportunities to develop new approaches to pest management. Aggregation pheromones and plant-derived semiochemicals are already available in commercial products. We review these semiochemicals and consider how we can move away from using them mainly for monitoring to using them for control. We still know very little about the behavioral responses of thrips to semiochemicals, and we show that research in this area is needed to improve the use of semiochemicals in pest management. We also propose that thrips should be used as a model system for semiochemically mediated behaviors of small insects that have limited ability to fly upwind.

Oviposition preference and olfactory response of Diaphania indica (Lepidoptera: Pyralidae) to volatiles of uninfested and infested cucurbitaceous host plants

The cucumber moth, Diaphania indica (Saunders) (Lepidoptera: Pyralidae), is a major pest of cucurbitaceous plants. The oviposition preference and olfactory response of larvae, mated and unmated male and female adults to volatiles emanating from uninfested and infested plants of four species of cucurbitaceous host plants and odours of conspecifics were recorded. Also the role of experience in the host finding behaviour of D. indica was evaluated. The experiments were done using a wind tunnel, olfactometer attraction assays and oviposition bioassays. The results reveal that fewer eggs were laid on infested plants than on uninfested plants. Females significantly preferred cucumber over squash, melon and watermelon. Cucurbitaceous plants elicited adults of D. indica to fly upwind followed by landing on the plants. The effect of experience on the olfactory preference of D. indica was dependent on the host plant. Females that had experience of cucumber, squash and melon plants were significantly attracted to the same plant, but not in the case of watermelon. Larvae of this pest were attracted only to volatiles of uninfested cucumber, squash and melon, whereas volatiles of conspecifics, infested plants and intact watermelon did not attract larvae. This study is an initial attempt to investigate the role of volatile infochemicals in the host-finding behaviour of D. indica. These results provide baseline information for the development of new control strategies against D. indica.

Mate-location flight of the red-necked longicorn beetle, Aromia bungii (Coleoptera: Cerambycidae): an invasive pest lethal to Rosaceae trees

The red-necked longicorn beetle, Aromia bungii (Faldermann) (Coleoptera: Cerambycidae), which has recently invaded Japan, is a serious pest species. The larvae kill cherry and orchard Rosaceae trees, such as peach, plum, and apricot. To clarify their features for mate location, male and female flight behaviors were observed in wind tunnels with caged males and females as lures. In a small wind tunnel (50 cm in diameter, 2 m in length), both sexes showed takeoff behavior according to increasing airflow. The rate of female takeoff against the male lure, a live male in a wire-netting cage, tended to be higher than for other combinations. In a large wind tunnel (ca. 1.6 m in diameter, 4 m in length), both sexes ascended and showed momentary hovering and astatic flight. When male lure cages were placed windward in the wind tunnel, females showed “upwind flight,” while males did not. It is assumed that females were induced to fly upwind by a factor derived from live males, likely to be an airborne pheromone component.

Bumblebees minimize control challenges by combining active and passive modes in unsteady winds

The natural wind environment that volant insects encounter is unsteady and highly complex, posing significant flight-control and stability challenges. It is critical to understand the strategies insects employ to safely navigate in natural environments. We combined experiments on free flying bumblebees with high-fidelity numerical simulations and lower-order modeling to identify the mechanics that mediate insect flight in unsteady winds. We trained bumblebees to fly upwind towards an artificial flower in a wind tunnel under steady wind and in a von Kármán street formed in the wake of a cylinder. Analysis revealed that at lower frequencies in both steady and unsteady winds the bees mediated lateral movement with body roll - typical casting motion. Numerical simulations of a bumblebee in similar conditions permitted the separation of the passive and active components of the flight trajectories. Consequently, we derived simple mathematical models that describe these two motion components. Comparison between the free-flying live and modeled bees revealed a novel mechanism that enables bees to passively ride out high-frequency perturbations while performing active maneuvers at lower frequencies. The capacity of maintaining stability by combining passive and active modes at different timescales provides a viable means for animals and machines to tackle the challenges posed by complex airflows.

Flight Tunnel Response of Male European Corn Borer Moths to Cross-Specific Mixtures of European and Asian Corn Borer Sex Pheromones: Evidence Supporting a Critical Stage in Evolution of a New Communication System.

Previous flight tunnel studies showed that 3-5 % of male European corn borer (ECB) moths, Ostrinia nubilalis, could fly upwind and make contact with sources releasing the sex pheromone of the closely related Asian corn borer (ACB), Ostrina furnacalis, [2:1 (Z)-12-tetradecenyl acetate (Z12-14:OAc) : (E)-12-teradecenyl acetate (E12-14:OAc)] and that 2-4 % of ACB males could similarly fly upwind to the sex pheromone blends of the ECB Z- [97:3 (Z)-tetradecenyl acetate (Z11-14:OAc) : (E)-tetradecenyl acetate (E11-14:Ac)] and E-strains (1:99 Z/E11-14:OAc) pheromones. The results supported the hypothesis that the evolution of the ACB pheromone system from an ECB-like ancestor included a stage in which males could be attracted to the unusual females emitting Z12- and E12-14:OAc while retaining their responsiveness to the ancestral pheromone blend of Z11- and E11-14:OAc. Here, we showed further that ECB E-strain males exhibited upwind oriented flight and source contacts to sources containing all combinations of ECB and ACB components. Maximal response levels were observed with the E-strain 99:1 E11/Z11-14:OAc blend, and high response levels also were observed with two other blends containing E11-14:OAc as the major component (E11:E12 and E11:Z12). Upwind flight and source contact also occurred at lower levels with the remaining blend combinations in which Z11-, E12-, or Z12-14:OAc was the major component. Our current results support the hypothesis concerning the evolution of ACB from an ECB-like ancester by showing that males were able to respond to females producing either the 12-14:Ac isomers, 11-14:Ac isomers, or even mixtures of all four components.

Towards an Understanding of Molecule Capture by the Antennae of Male Beetles Belonging to the Genus Rhipicera (Coleoptera, Rhipiceridae).

Working on the hypothesis that an important function of the lamellate antennae of adult male beetles belonging to the genus Rhipicera is to detect scent associated with female conspecifics, and using field observations, anatomical models derived from X-ray microcomputed tomography, and scanning electron microscopy, we have investigated the behavioral, morphological, and morphometric factors that may influence molecule capture by these antennae. We found that male beetles fly upwind in a zigzag manner, or face upwind when perching, behavior consistent with an animal that is tracking scent. Furthermore, the ultrastructure of the male and female antennae, like their gross morphology, is sexually dimorphic, with male antennae possessing many more of a particular type of receptor-the sensillum placodeum-than their female counterparts (approximately 30,000 vs. 100 per antenna, respectively). Based on this disparity, we assume that the sensilla placodea on the male antennae are responsible for detecting scent associated with female Rhipicera beetles. Molecule capture by male antennae in their alert, fanned states is likely to be favoured by: (a) male beetles adopting prominent, upright positions on high points when searching for scent; (b) the partitioning of antennae into many small segments; (c) antennal morphometry (height, width, outline area, total surface area, leakiness, and narrow channels); (d) the location of the sensilla placodea where they are most likely to encounter odorant molecules; and (e) well dispersed sensilla placodea. The molecule-capturing ability of male Rhipicera antennae may be similar to that of the pectinate antennae of certain male moths.

Volatile semiochemical-conditioned attraction of the male yellow fever mosquito, Aedes aegypti, to human hosts.

We investigated the olfactory responses of male mosquitoes to kairomones of vertebrate hosts in a dual-port olfactometer. The behavioral responses of unmated and mated male and female mosquitoes from one to ten days old to human odors were compared to the odors of different human hosts. To evaluate the relationship between the age of male mosquitoes and their responses, we performed experiments with males at different ages. Unmated Ae. aegypti males, one to two days old, did not fly upwind to human odor, whereas between three and ten days old they exhibited increased flight activity. The results showed that unmated and mated females were attracted by human odor, but those mated were more attracted by human odor than when unmated. Mated males were, in general, attracted by human odor, while the unmated males were not attracted but showed increased flight activity in the presence of human odor, suggesting swarming behavior. Further studies should be carried out in order to determine the role of human odors on male Ae. aegypti behavior.

Pheromone modulates plant odor responses in the antennal lobe of a moth.

In nature, male moths are exposed to a complex plant odorant environment when they fly upwind to a sex pheromone source in their search for mates. Plant odors have been shown to affect responses to pheromone at various levels but how does pheromone affects plant odor perception? We recorded responses from neurons within the non-pheromonal "ordinary glome ruli" of the primary olfactory center, the antennal lobe (AL), to single and pulsed stimulations with the plant odorant heptanal, the pheromone, and their mixture in the male moth Agrotis ipsilon. We identified 3 physiological types of neurons according to their activity patterns combining excitatory and inhibitory phases. Both local and projection neurons were identified in each physiological type. Neurons with excitatory responses to heptanal responded also frequently to the pheromone and showed additive responses to the mixture. Moreover, the neuron's ability of resolving successive pulses generally improved with the mixture. Only some neurons with combined excitatory/inhibitory, or purely inhibitory responses to heptanal, also responded to the pheromone. Although individual mixture responses were not significantly different from heptanal responses in these neurons, pulse resolution was improved with the mixture as compared with heptanal alone. These results demonstrate that the pheromone and the general odorant subsystems interact more intensely in the moth AL than previously thought.

Age-based mating success in the codling moth,Cydia pomonella, and the obliquebanded leafroller,Choristoneura rosaceana

In this study, the passage of spermatophores was examined between 1-day-old males mated in no-choice situations with females 0, 2, 4, or 6 days old and the converse for both the codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), and the obliquebanded leafroller, Choristoneura rosaceana (Harris). For C. pomonella, female age had no effect on the passage of spermatophores from males, and only 6-day-old female C. rosaceana had reduced spermatophore number. The ages of moths at the time of mating had a greater effect on males, with C. pomonella males older than 2 days showing significant reductions in the ability to successfully pass a spermatophore to 1-day-old females. For C. rosaceana, 2- and 6-day-old males were significantly less likely to pass a spermatophore, but reduced transfer from 4-day-old males did not reach statistical significance. Wind-tunnel assays were used to evaluate the ability of 1- or 4-day-old males to fly upwind and successfully contact a young calling female. Four-day-old males were more likely to initiate flight upwind, but were less efficient at finding and contacting the females than younger males. This study suggests that evaluation of multiple components of the mating process are required to understand the effect of age at the time of mating on population dynamics of these moths.

The role of vision in plume tracking by flying Drosophila

Event Abstract Back to Event The role of vision in plume tracking by flying Drosophila Michael H. Dickinson1* and Floris Van Breugel2 1 University of Washington, Biology, United States 2 Caltech, CDS, United States Finding food is one of the most critical tasks animals perform on a routine basis. Many animals accomplish this by following attractive odor plumes back to their source, which is a complex problem involving the integration of multiple sensory modalities. A common iterative strategy observed in many flying animals is to surge upwind when detecting an odor filament, and then cast crosswind when losing the odor. Both components of this behavior are enigmatic, because animals do not have a direct measure of either their groundspeed or the wind velocity. Nevertheless, animals are able to fly upwind and crosswind across a range of flight velocities, wind speeds, and body orientations. Despite a century of quantitative scientific research on this subject, we still do not fully understand how animals track plumes. To address this question we focused on the fruit fly, Drosophila, because its small size and robust indoor behavior makes it possible to observe long and uninterrupted bouts of odor tracking in the presence of an structured odor plume. Using a novel experimental setup we can estimate a fly's visual and olfactory experience throughout a long flight trajectory, and by automating the data collection we have analyzed over 10,000 flight trajectories under a variety of experimental conditions. By projecting different patterns on the walls of the wind tunnel we can directly manipulate the visual cues available during different components of the behavior. We can also alter mechanosensory and olfactory cues by manipulating the antennae of the flies being tested. Using the large data set we show how flies use vision, olfaction, and wind sensing to follow an attractive odor plume to its source. Within 200 milliseconds of detecting the plume odor, the flies turn upwind (i.e. surge) using visual cues. 400-800 milliseconds after losing the plume, they initiate a crosswind cast, which relies on both visual and antennal cues. This is a robust behavior across a range of wind speeds, indicating that flies must have some ability by which they estimate wind speed and direction. Using tools from nonlinear control theory we have developed a model of how flies accomplish this task. Our results have implications for the underlying neural implementation of odor tracking behavior, as well as more general issues relate to the sensory control of flight control. Acknowledgements This research was supported by grants from the Air Force Office of Scientific Research (AFOSR) to MHD, a Hertz Fellowship and a National Science Foundation (NSF) Graduate Research Fellowship to FvB. Keywords: Flight control, plume tracking, Control theory, Drosophila melanogaster, Vision Conference: International Conference on Invertebrate Vision, Fjälkinge, Sweden, 1 Aug - 8 Aug, 2013. Presentation Type: Oral presentation preferred Topic: The visual control of flight and locomotion Citation: Dickinson MH and Van Breugel F (2019). The role of vision in plume tracking by flying Drosophila. Front. Physiol. Conference Abstract: International Conference on Invertebrate Vision. doi: 10.3389/conf.fphys.2013.25.00039 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 28 Feb 2013; Published Online: 09 Dec 2019. * Correspondence: Dr. Michael H Dickinson, University of Washington, Biology, Seattle, United States, flyman5@gmail.com Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Michael H Dickinson Floris Van Breugel Google Michael H Dickinson Floris Van Breugel Google Scholar Michael H Dickinson Floris Van Breugel PubMed Michael H Dickinson Floris Van Breugel Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Sexual Behavior of <I>Grapholita molesta</I> and <I>Choristoneura rosaceana</I> (Lepidoptera: Tortricidae) in a Flight Tunnel After Prolonged Exposure to the Aerial Concentration of Pheromone Previously Measured in Orchards Treated With Pheromone for Mating Disruption

Sexual behavior of male moths after prolonged exposure to the 1-ng pheromone/m(3) air previously measured in orchards treated with pheromone for mating disruption was examined in a flight tunnel. The exposure of Grapholita molesta (Busck) to 1-ng (Z)-8-dodecen-1-yl acetate (Z8-12:OAc)/m(3) air for 15 min had no effect on their ability to fly upwind to a conspecific, virgin calling female. After 30 min of exposure, males exposed to a control treatment were 1.4× more likely orient to a female than males exposed to pheromone-treated air. Some G. molesta males retained the ability to orient to a female after a 30-min exposure period when the aerial concentration of Z8-12:OAc was increased 500,000× to 0.5 gm/m(3). Prolonged exposure to Z8-12:OAc did not affect response to a synthetic pheromone lure. The time required to initiate behavioral responses to a female or a lure was not affected by pheromone exposure. Male Choristoneura rosaceana (Harris) exposed to a control treatment for 15 min were 38.5× more likely to orient to a conspecific, virgin calling female than males exposed to 1-ng (Z)-11-tetradecen-1-yl acetate (Z11-14:OAc)/m(3) air for 15 min. After 30 min of exposure males were unable to fly upwind to a female. Males exposed to a control treatment for 15 min were 4.3× more likely to fly upwind to a synthetic pheromone lure than males exposed to 1-ng Z11-14:OAc/m(3) air for 15 min. The time required to initiate behavioral responses to a female or a lure was not affected by exposure to pheromone.

Wind direction and proximity to larval sites determines malaria risk in Kilifi District in Kenya.

Studies of the fine-scale spatial epidemiology of malaria consistently identify malaria hotspots, comprising clusters of homesteads at high transmission intensity. These hotspots sustain transmission, and may be targeted by malaria-control programmes. Here we describe the spatial relationship between the location of Anopheles larval sites and human malaria infection in a cohort study of 642 children, aged 1–10-years-old. Our data suggest that proximity to larval sites predict human malaria infection, when homesteads are upwind of larval sites, but not when homesteads are downwind of larval sites. We conclude that following oviposition, female Anophelines fly upwind in search for human hosts and, thus, malaria transmission may be disrupted by targeting vector larval sites in close proximity, and downwind to malaria hotspots.

Altered Olfactory Receptor Neuron Responsiveness Is Correlated with a Shift in Behavioral Response in an Evolved Colony of the Cabbage Looper Moth, Trichoplusia ni

There is little understanding of how sex pheromone blends might change during speciation events. For the cabbage looper, Trichoplusia ni, there is a mutant laboratory strain that has exhibited characteristics of a shift to a new pheromone blend. Mutant females produce a blend that is significantly different from wild-type females in having a much higher proportion of a minor pheromone component and lower quantity of the major component. Males in this colony have changed over the years to become more broadly tuned and fly upwind equally well to both the wild-type and mutant female pheromone blends. They also exhibit reduced overall sensitivity to pheromone, flying upwind to either blend at a lower success rate than is typical when wild-type males respond to the wild-type blend. Using single-cell recordings, we examined the olfactory receptor neurons (ORNs) of males from evolved and wild-type colonies for evidence of changes in response characteristics that might explain the above-described behavioral evolution. We found that in evolved-colony males the ORNs tuned to the major sex pheromone component exhibited a somewhat lower responsiveness to that compound than the ORNs of wild-type males. In addition, the minor pheromone component, emitted at excessively high rates by mutant females, elicited a drastically reduced ORN responsiveness in evolved-colony males compared to wild-type males. This alteration in ORN responsiveness may be responsible for allowing evolved males to tolerate the excessive amounts of the minor pheromone component in the mutant female blend, which would normally antagonize the upwind flight of unevolved males. Thus, peripheral olfactory alterations have occurred in T. ni males that are correlated with the evolution of the more broadly tuned, but less sensitive, behavioral response profile.

Olfactory neuron responsiveness and pheromone blend preference in hybrids between Ostrinia furnacalis and Ostrinia nubilalis (Lepidoptera: Crambidae)

The olfactory receptor neuron (ORN) and behavioral responses of hybrids between the Asian corn borer (ACB), Ostrinia furnacalis, and the E-strain European corn borer (ECB(E)), Ostrinia nubilalis were examined and compared to the parental populations. In hybrids and both parents, the large-spike-size ORN was capable of responding to all four pheromone components of ACB and ECB, despite differences in which compounds elicited the greatest spike frequency in each population. There was a small-spiking ORN more narrowly tuned to the minor pheromone components in both ACB and ECB(E). In hybrids the homologous small-spiking ORN was tuned primarily to the ECB(E) minor pheromone component, with some responsiveness to the ACB minor component. Both species and all the hybrids had an intermediate spike-size ORN tuned primarily to their common behavioral antagonist. Dominance of responsiveness to the ECB(E) versus the ACB minor pheromone component on the small-spiking ORN may explain the greater tendency of hybrids to fly upwind to the ECB(E) pheromone blend than the ACB blend. This finding points toward a distinct evolutionary role for this ORN in allowing a pheromone shift.

A search for components in human body odour that attract females of Aedes aegypti.

In a new type of wind tunnel, mosquitoes fly upwind towards host odour sources and towards human skin wash extracts obtained by rubbing the skin with a pad soaked in ethanol. We used this behavioural response as a bioassay to identify attractants in liquid chromatography fractions of such extracts. L-Lactic acid is a major constituent of skin wash extracts and it is a necessary component for the extract's effectiveness. As a single stimulus, however, L-lactic acid is only slightly effective. This indicates that the extract's high degree of effectiveness is based on a synergism of L-lactic acid and other odour components. The separation of the extract by liquid chromatography revealed three distinct regions of active fractions, only one of which contained L-lactic acid. The components of the other two regions have not yet been determined. A combination of fractions in these two regions together with L-lactic acid is as attractive as the complete extract.