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We characterized single upwind surges of flying male Helothis virescens moths in response to individual strands of pheromone generated experimentally in a wind tunnel. We then showed how this surge functions in this species as a basic 13.4-cm, 0.38-sec-long building block that is strung together repeatedly during typical male upwind flight in a normal pheromone plume. The template for a single iteration, complete with crosswind casting both before and after the straighter upwind surging portion, was exhibited by males flying upwind to pheromone and experiencing ifiament contactsjust frequently enough to produce successful upwind flight to the source, as hypothesized by an earlier model. Also as predicted, with more frequent filament contact by males, only the straightest upwind portions of the surges were reiterated, producing direct upwind flight with little crosswind casting. Electroantennogram recordings made from males in free Tfight upwind in a normal point source pheromone plume further support the idea that a high frequency offilaments encountered under the usual pheromone plume conditions promotes only these repeated straight surges. In-flight electroantennogram recordings also showed that when filament contacts cease, the casting, counterturning program begins to be expressed after a latency period of 0.30 sec. Together these results provide a plausible explanation for how male and female moths, and maybe other insects, fly successlly upwind in an odor plume and locate the source of odor, using a surging-casting, phasictonic response to the onset and disappearance of each odor strand.In the quest for understanding how male moths fly upwind and locate females (1-5) there have been suggestions (6, 7) that all odor-mediated flight in moths, including host plant location by females, may be explained by two programs, optomotor anemotaxis (2) and self-steered counterturning (8), that are turned on and modulated by odor fluctuations. It has also been pointed out (9, 10) that many other kinds of insects flying upwind to odor exhibit behavior somewhat similar to moths', and these other insects may also use these same mechanisms in odor-source location. Knowledge about the mechanisms used by moths should therefore be important for understanding the neuroethology of olfaction, the evolution of pheromone and host-plant-insect systems, and the potential application of pheromones in insect control.The physical structure of a pheromone plume created by a point source of odor is not a time-averaged homogeneous cloud. Turbulence causes the plume to break up into strands of odor-laden air (filaments) interspersed with pockets of clean air where little or no odor is present (11,12). The physical intermittency of these filaments is a requirement for male moths to sustain their upwind progress; they will not continue to fly upwind upon entering a homogeneous cloud of pheromone (4, 13) but will if this cloud is alternated with swaths of clean air (14).Recently, results from experiments investigating the highspeed beh...

Insect olfactory systems present models to study interactions between animal genomes and the environment. They have evolved for fast processing of specific odorant blends and for general chemical monitoring. Here, we review molecular and physiological mechanisms in the context of the ecology of chemical signals. Different classes of olfactory receptor neurons (ORNs) detect volatile chemicals with various degrees of specialization. Their sensitivities are determined by an insect-specific family of receptor genes along with other accessory proteins. Whereas moth pheromones are detected by highly specialized neurons, many insects share sensitivities to chemical signals from microbial processes and plant secondary metabolism. We promote a more integrated research approach that links molecular physiology of receptor neurons to the ecology of odorants.

The neural computations used to represent olfactory information in the brain have long been investigated. Recent studies in the insect antennal lobe suggest that precise temporal and/or spatial patterns of activity underlie the recognition and discrimination of different odours, and that these patterns may be strengthened by associative learning. It remains unknown, however, whether these activity patterns persist when odour intensity varies rapidly and unpredictably, as often occurs in nature. Here we show that with naturally intermittent odour stimulation, spike patterns recorded from moth antennal-lobe output neurons varied predictably with the fine-scale temporal dynamics and intensity of the odour. These data support the hypothesis that olfactory circuits compensate for contextual variations in the stimulus pattern with high temporal precision. The timing of output neuron activity is constantly modulated to reflect ongoing changes in stimulus intensity and dynamics that occur on a millisecond timescale.

Previous studies have suggested that Plasmodium parasites can manipulate mosquito feeding behaviours such as probing, persistence and engorgement rate in order to enhance transmission success. Here, we broaden analysis of this ‘manipulation phenotype’ to consider proximate foraging behaviours, including responsiveness to host odours and host location. Using Anopheles stephensi and Plasmodium yoelii as a model system, we demonstrate that mosquitoes with early stage infections (i.e. non-infectious oocysts) exhibit reduced attraction to a human host, whereas those with late-stage infections (i.e. infectious sporozoites) exhibit increased attraction. These stage-specific changes in behaviour were paralleled by changes in the responsiveness of mosquito odourant receptors, providing a possible neurophysiological mechanism for the responses. However, we also found that both the behavioural and neurophysiological changes could be generated by immune challenge with heat-killed Escherichia coli and were thus not tied explicitly to the presence of malaria parasites. Our results support the hypothesis that the feeding behaviour of female mosquitoes is altered by Plasmodium, but question the extent to which this is owing to active manipulation by malaria parasites of host behaviour.

Single-cell electrophysiological recordings were obtained from olfactory receptor neurons in sensilla trichodea on male antennae of the heliothine species Heliothis subflexa and the closely related congener H. virescens. A large percentage of sensilla (72% and 81%, respectively, of all sensilla sampled) contained a single odor-responsive receptor neuron tuned to the major pheromone component of both species, Z-11-hexadecenal. A second population of sensilla on H. subflexa antennae (18%) housed receptor neurons that were tuned to Z-9-hexadecenal but also responded with less sensitivity to Z-9-tetradecenal. A similar population of sensilla (4%) on H. virescens male antennae housed receptor neurons that were shown to be tuned specifically only to Z-9-tetradecenal, with no response to even high dosages of Z-9-hexadecenal. A third population of sensilla (comprising 8% and 16% of the sensilla sampled in H. subflexa and H. virescens, respectively) housed two olfactory receptor neurons, one of which was tuned to Z-11-hexadecenyl acetate and the other tuned to Z-11-hexadecenol. In H. subflexa the Z-11-hexadecenyl acetate-tuned neuron also responded to Z-9-tetradecenal with nearly equivalent sensitivity. The behavioral requirements of males of these two species for distinct pheromonal blends was, therefore, reflected by the subtle differences in the tuning properties of antennal olfactory receptor neurons.

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