Moth sex pheromone adsorption to leaf surface: bridge in time for chemical spies

Abstract When brussels sprouts leaves were exposed to air passed over a single calling Mamestra brassicae moth, sex pheromone was adsorbed onto the leaf surface to such an extent that it subsequently elicited behavioural responses in conspecific male moths, as well as in female Trichogramma evanescens , egg parasitoids of M. brassicae. Male moths responded to odour-treated leaves over a short distance (c. 5 cm) in a wind tunnel, but were not attracted at a distance of 1 m. In contrast, sex pheromone adsorbed to and re-released from the glass wall of a wind tunnel compartment attracted male moths from 1 m. Trichogramma's locomotory behaviour on leaves is described. On leaves previously exposed to air passed over a calling female moth, Trichogramma wasps stayed significantly longer on leaves treated with clean air or air passed over a non-calling female moth. In addition, wasps spent relatively more time along the margin and on the leaf underside on treated leaves compared with control leaves. All effects persisted for at least 4 h, and after 24 h treated leaves still increased wasp residence times. These results are the first example of responses of male moths to adsorbed airborne sex pheromone originating from a single female moth. They further suggest how sex pheromone released by nocturnal moths may function as a kairomone for diurnally foraging parasitoids.     

Odour-conditioned anemotaxis of apterous aphids (Cryptomyzus korschelti) in response to host plants

ABSTRACT. Orientation responses of adult apterous virginoparae of Cryptomyzus korschelti Börner were recorded using a locomotion-compensator in front of a wind tunnel. Individual aphids were tested under four consecutive treatments: without wind; clean wind; and wind carrying odour of the host plant Stachys sylvatica or odour of a non-host plant Solanum tuberosum. The walking tracks were tortuous in all treatments except when the odour of host plants was used. Host plant odour induced upwind orientation of aphids (odour-conditioned positive anemotaxis). Track variables such as vector length, straightness, upwind time and upwind length, increased when the aphids moved upwind. Walking speeds were not affected. The simultaneous stimulation by wind and host plant odour caused aphids to walk upwind for more than 1 m in 10 min. These findings suggest that olfactory attraction of aphids is involved in host plant selection.     

Odor specificity testing in the assessment of efficacy and non-target risk for Trichogramma ostriniae (Hymenoptera: Trichogrammatidae)

Trichogramma ostriniae Pang and Chen (Hymenoptera: Trichogrammatidae) is an indigenous egg parasitoid of Ostrinia furnacalis Guenée (Lepidoptera: Crambidae) in China. We evaluated T. ostriniae's responses in olfactometer and wind tunnel assays to various host and plant odors that are likely to impact the efficacy and non-target risk of utilizing T. ostriniae as an augmentative biocontrol agent against Ostrinia nubilalis Hübner in the USA. In a Y-olfactometer, female T. ostriniae exhibited innate positive responses to the egg mass volatiles, scale volatiles, and synthetic sex pheromones of O. nubilalis. When exposed to O. nubilalis pheromone while walking on a platform in a wind tunnel, the wasps manifested significant changes in patch exploration behavior, including delayed dispersal from the platform and slowed walking. The wasps did not respond innately to the synthetic pheromone of a non-target species, Spodoptera frugiperda J. E. Smith (Lepidoptera: Noctuidae), however. Exposing wasps to S. frugiperda pheromone together with eggs of a factitious rearing host prior to testing also did not alter the wasps’ lack of response to the pheromone, indicating that associative learning of the novel odor did not occur. Lastly, wasps showed no innate responses to leaf volatiles from corn (Zea mays L.) and pepper (Capsicum annuum L.), two crops attacked by O. nubilalis. We conclude that T. ostriniae is likely to be highly efficient at finding O. nubilalis eggs in the field through the exploitation of host chemical cues. Further, T. ostriniae's response to moth pheromone appears to be relatively host-specific, since the wasps responded to the pheromone of a congener to their natural host, but not to the pheromone of a more distantly related non-target species. This type of odor-specificity could be an important mechanism for reducing the risk of T. ostriniae attack on non-target species.     

Oriented responses of the triatomine bugs Rhodnius prolixus and Triatoma infestans to vertebrate odours on a servosphere

Oriented responses of both R. prolixus and T. infestans adults were recorded on a servosphere to mouse-odour, one of its components (CO₂), and to rabbit urine-odour. The volatiles were delivered in an air-stream under controlled conditions which excluded other sensory modalities. In stimulus-free air the triatomines walked preferentially downwind in straight bouts interrupted by stops or periods at relatively low speeds, all of variable duration. In odour-laden air, bugs maintained their typical walking habit but switched from negative to positive anemotaxis. The characteristic response to odour onset was to stop, sample the air with the antennae, turn upwind in situ, and then walk off in the direction of the source for at least a few seconds, i.e., odour mediated anemotaxis. Mouse-odour caused T. infestans to increase its speed to 5.3 cms^-1. Both species continued with the upwind response for some time after odour delivery ceased, but the crosswind component of the tracks was more prominent during this period — an effort, we presume, by the bugs to re-contact an odour plume. This investigation provides unequivocal evidence for host finding in triatomines by olfactory cues alone.     

Identification of compounds from the eggs of Ostrinia nubilalis and Mamestra brassicae having kairomone activity on Trichogramma brassicae

In the presence of an extract of Ostrinia nubilalis or Mamestra brassicae eggs, female Trichogramma brassicae exhibited increased rates of upwind locomotion in the tubes of a linear olfactometer. GC and GC-MS analyses of O. nubilalis and M. brassicae egg extracts revealed the presence of fatty acids, their ethyl esters, and various hydrocarbons. Exposing the wasps to a mixture of the five main saturated hydrocarbons (heneicosane, tricosane, pentacosane, heptacosane and nonacosane) increased the upwind progression in the olfactometer. Single hydrocarbons elicited reduced or no activity. Ethyl palmitate and palmitic acid were also effective in increasing upwind locomotion. Z11-14:Ac, the main component of the female sex pheromone of O. nubilalis was inactive, regardless of its concentration. It is concluded that various compounds present on the surface of the O. nubilalis egg masses may play a role in the orientation of T. brassicae to its host.     

Flight behaviour of the cabbage seedpod weevil

The influence of host odour, windspeed, position of the sun, and temperature on flight behaviour of the cabbage seedpod weevil (Ceutorhynchus assimilis Paykull) were studied. This weevil showed a positive anemotaxis (upwind flight) inside the odour plume of a host crop (Brassica napus L.). Outside the odour plume the weevil showed a pronounced phototaxis at windspeeds below 1.5 m/s. At higher windspeeds, the seedpod weevils flew downwind. The cabbage seedpod weevil flies most readily at low windspeeds (less than 0.5 m/s) and at temperatures above 22 °C.     

繁育寄主对三种赤眼蜂个体大小及抱卵量的影响

【目的】繁育寄主及成蜂日龄均会影响赤眼蜂体内的抱卵量,为了解繁育寄主及育出赤眼蜂个体大小与赤眼蜂抱卵量的关系,我们在室内研究了米蛾 Corcyra cephalonica（Stainton）卵和亚洲玉米螟 Ostrinia furnacalis （Güenée）卵作为繁育寄主对3种赤眼蜂雌蜂大小、抱卵量的影响。【方法】在光周期14L:10D、温度25±1℃的室内条件下,以米蛾卵、亚洲玉米螟卵作为繁育寄主,解剖不同日龄（羽化后12-72 h）松毛虫赤眼蜂 Trichogramma dendrolimi Matsumura、玉米螟赤眼蜂Trichogramma ostriniae Pang et Chen以及稻螟赤眼蜂Trichogramma japonicum Ashmead,观察雌蜂抱卵量,测量成蜂的大小,分析雌蜂抱卵量与其日龄、个体大小以及繁育寄主的关系【结果】在羽化后48 h以内,3种蜂的抱卵量随雌蜂日龄的增长而增加。羽化后72 h,除米蛾卵繁育的玉米螟赤眼蜂及玉米螟卵繁育的松毛虫赤眼蜂的抱卵量比羽化后48 h时略高外,其他组合均出现下降。以亚洲玉米螟卵为繁育寄主时的玉米螟赤眼蜂的抱卵量显著高于对应日龄的以米蛾卵为繁育寄主时的抱卵量,而松毛虫赤眼蜂（除羽化后12和24 h外）和稻螟赤眼蜂则相反;以米蛾卵为繁育寄主时,相同日龄的赤眼蜂中,松毛虫赤眼蜂抱卵量最高,稻螟赤眼蜂的抱卵量最低;以亚洲玉米螟卵为繁育寄主时,玉米螟赤眼蜂抱卵量最高,稻螟赤眼蜂抱卵量最低。雌蜂抱卵量与雌蜂个体大小呈线性正相关,相关程度随雌蜂日龄增加而更加明显。【结论】后足胫节长度可以作为评价松毛虫赤眼蜂、玉米螟赤眼蜂和稻螟赤眼蜂质量的指标。赤眼蜂抱卵量受繁育寄主、赤眼蜂蜂种及雌蜂日龄的显著影响,寄主的适合性也会影响育出赤眼蜂的大小,筛选合适的繁育寄主有助于提高赤眼蜂的质量。     

Evidence of active or passive downwind dispersal in mark–release–recapture of moths

Modelling moth dispersal in relation to wind direction and strength could greatly enhance the role of pheromone traps in biosecurity and pest management applications. Anemotaxis theory, which describes moth behaviour in the presence of a pheromone plume and is used as a framework for such models. Currently, however, that theory includes only three components: upwind, zigzagging, and sideways casting behaviour. We test anemotaxis theory by analysing the data from a series of mark–release–recapture experiments where the wind direction was known and the insects were trapped using an irregular grid of pheromone traps. The trapping results provide evidence of a downwind component to the flight patterns of the released insects. This active or passive downwind dispersal is likely to be an appetitive behaviour, occurring prior to the elicitation of pheromone‐oriented flight patterns (pheromone anemotaxis). Given the potential for significant displacement during downwind dispersal, this component will have impact on final trap captures and should be considered when constructing moth dispersal models.     

Host instar, body size and fitness in the koinobiotic parasitoid Aphidius nigripes

In aphidiine parasitoids, resources for growth and adult body size increase with host instar used by ovipositing females, but the fitness consequences of body size on fitness are poorly documented. We compared the fitness of male and female A. nigripesadults that varied in size as a consequence of developing in different instars of their host Macrosiphum euphorbiae. When reproductive fitness was measured without considering time, female wasps from small and large hosts performed similarly, contributing 125–175 foundresses plus 100–180 sons to the next generation. However, when expressed as the innate capacity for increase (r _m), female fitness correlated with host-induced variation of wasp size, indicating that micropopulations initiated by large wasps would increase faster. In a wind-tunnel, a sex pheromone plume from large female wasps induced more males to fly upwind when released at a distance of 50 cm downwind than small females, indicating that large females were sexually more attractive. With respect to male body size effects on fitness, large individuals performed similar to small ones, whether fitness was measured by lifetime mating frequency, fertile inseminations, or proportion of daughters among progeny born from their mates. When young naive males of unequal size were directly competing for mating with a virgin female, small and large males had equal mating success, and large individuals were no more successful than small ones at displacing a competitor already positioned on a receptive female. In a wind-tunnel test where males were scored on their ability to reach a female pheromone source, small and large males were equally affected by wind speed but reached the source located 50 cm downwind in equal proportions, suggesting similar capacity for finding mates by flying upwind. Our results indicate that despite host resources not being fixed at the time of attack for the koinobiont A. nigripes, fitness consequences of resource limitation by the mother may be perceived to be greater for daughters than sons, which would explain male-biased sex ratio in early-instar hosts.     

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.     

Differential response of Trichogramma wasps to extreme sex pheromone types of the noctuid moth Heliothis virescens

1. Chemical espionage in nature may occur when predators or parasitoids home in on animal or plant communication signals. Parasitoid wasps are known to use pheromones emitted by adults hosts to locate host eggs, larvae or pupae. The response of Trichogramma egg parasitoids to a synthetic sex pheromone blend of moths has been shown in a number of studies over the past 40 years. 2. Trichogramma pretiosum (Hymenoptera, Trichogrammatidae) is a tiny parasitic wasp, attacking the eggs of the noctuid moth Heliothis virescens (Lepidoptera, Noctuidae). This study investigated whether T. pretiosum homes in on the sex pheromone of H. virescens at close range. The arrestment response of the wasps to sex pheromone gland extracts of two types of female moths, so‐called high and low females, was also tested, referring to two selected extreme pheromone types of H. virescens. The study also investigated whether the wasps would mount females, possibly to hitchhike with them. 3. The wasps were arrested by the common, ‘low’ pheromone, but not by the rare, ‘high’ pheromone or by extracts from male hairpencils. The wasps did not show a preference for separate sex pheromone compounds, but when pre‐exposed to the major sex pheromone component of H. virescens before the tests together with H. virescens eggs, they did show a preference, indicating learning behaviour. In the mounting experiments, mated females were mounted significantly more than virgin females or males, suggesting that hitchhiking is a strategy used by these wasps to locate moth eggs. 4. This represents the first study to show a differential response of parasitoid wasps to two different sex pheromone types in a single host species. The results warrant further investigations into the potential role of parasitic wasps in the evolution of sexual communication in moths.     

Guidance of flying male moths by wind-borne sex pheromone

ABSTRACT. On passing from clean air into a homogeneous cloud of sex pheromone in a wind tunnel flying male Adoxophyes orana (F.v.R.) (Lepidoptera: Tortricidae) turned more or less upwind and reduced the time and distance between their switchings of track from one side of the wind line to the other. These responses became adapted under the constant pheromone stimulation in the cloud, thereby arresting upwind progress; but the adapted moths would now 'lock-on' to an added pheromone plume and advance upwind along it. Moths also locked-on to the border of a pheromone cloud, not by turning back on losing the scent as previously supposed but by initiating the above programme of small-amplitude, crosswind movements (reversing anemomenotaxis). The onset and cessation of the pheromone stimulus produced anemotactic responses that differed quantitatively within a continuum, not two distinct kinds of response as previously supposed. The behavioural mechanism whereby uniform permeation of an area with synthetic sex pheromone can prevent males from finding females is reconsidered.     

Electroantennogram and behavioural responses of different forms of the bird cherry-oat aphid,Rhopalosiphum padi,to sex pheromone and a plant volatile

Winged (alate) virginoparae were induced in the laboratory by crowding the bird cherry-oat aphid, Rhopalosiphum padi (L.) under long-day conditions. Males and gynoparae (the winged female form that produces sexual females) were induced by short days. Electroantennogram (EAG) and behavioural responses were investigated in each of the three forms to two aphid sex pheromone components, (-)-(4aS,7S,7aR)-nepetalactol and (+)-(4aS,7S,7aR)-nepetalactone, and benzaldehyde, a volatile which is released by the winter host plant. All three compounds elicited EAG responses with the males showing the highest sensitivity to each compound. Both the nepetalactol and the nepetalactone elicited larger EAG responses in gynoparae than in the winged virginoparae but antennae from virginoparae were more responsive to benzaldehyde. Although the nepetalactone is not a sex pheromone component in R. padi the EAG responses were similar to those evoked by the nepetalactol, the sex pheromone, in all three aphid forms. In a linear-track olfactometer, significantly more male R. padi moved into air containing nepetalactol, nepetalactone or benzaldehyde than into a simultaneous choice of clean air (i.e. attraction) but nepetalactol was more attractive than nepetalactone. Males, however, showed no response to a mixture of nepetalactol and nepetalactone. Gynoparae were attracted only to the nepetalactol but were less sensitive than the males and showed no response to the nepetalactone or benzaldehyde. In contrast, alate virginoparae showed no behavioural responses to any of the compounds. The present study supports the idea that the male R. padi utilise both sex pheromone and benzaldehyde for mate/host-plant location in autumn. It also demonstrates, for the first time, polyphenic differences in the olfactory responses at the peripheral level between the two female forms. Such differences impact on the life-cycle strategy where winged virginoparae move between graminaceous summer host plants while gynoparae move from the summer hosts to the bird cherry, winter host. The latter move appears to be assisted by the sex pheromone released by sexual females, already present on that host, acting as an aggregation pheromone.     

逐日羽化的赤眼蜂品质的比较

本文报道了逐日羽化的赤眼蜂(Trichogramma maidis Pint.et Voeg)品质的比较结果.第1天羽化的雌蜂是群体中生活力最强的个体;第2和第3天羽化的雌蜂生殖力、寿命、个体大小和性比等4项指标逐渐下降;而第4天羽化的雌蜂质量最差.当逐日(第1—第4天)羽化的赤限蜂先分别用地中海粉斑螟卵繁殖1代,然后分别收集第1天内羽化的赤眼蜂试验时,子代(F₁)雌蜂的质量有了明显的提高.母代为第3和第4天羽化的赤限蜂,其子代雌蜂的质量分别超过母代.试验表明,通过实验室选择培养可以改进和提高大量培养赤眼蜂的质量.     

Kairomonal effect of sex pheromone components of two lepidopteran olive pests on Trichogramma wasps

Egg parasitoids are known to use a wide range of chemicals, emitted by plants, host eggs or adults, for host selection. The effect of the sex pheromone components of the lepidopteran olive pests Prays oleae (Lepidoptera: Yponomeutidae) and Palpita unionalis (Lepidoptera: Pyralidae) was studied under laboratory conditions, on the foraging behaviour of the egg parasitoid Trichogramma oleae (Hymenoptera: Trichogrammatidae). The response of T. oleae wasps to ( Z )-7-tetradecenal and ( E )-11-hexadecenal, major sex pheromone components of P. oleae and P. unionalis respectively, depended on the dose of the pheromone used in a Y-tube olfactometer bioassay. ( E )-11-hexadecenal elicited maximum attraction (70%) at a dose of 1 μg, while a dose of 100 μg ( Z )-7-tetradecenal attracted 80% of the tested wasps. ( E )-11-hexadecenyl acetate, the second sex pheromone component of P. unionalis , and the binary blend of ( E )-11-hexadecenyl acetate: ( E )-11-hexadecenal (7:3) were not attractive at these doses. The results of this research are discussed in view that they may be considered as alternatives in the biological control of these pests.     

Effects of intermittent and continuous pheromone stimulation on the flight behaviour of the oriental fruit moth, Grapholita molesta

ABSTRACT. When male oriental fruit moths, Grapholita molesta (Busck) (Tortricidae), casting in clean air entered an airstream permeated with pheromone their flight tracks changed immediately on initial contact with pheromone, but after a few seconds returned to casting as if in clean air. The degree of change in the flight track was directly related to the concentration of pheromone. Although little net uptunnel movement occurred in response to the continuous stimulation provided by a uniformly permeated airstream, when an intermittent stimulus provided by a point-source plume was superimposed onto the permeated airstream moths were able to 'lock on' and zigzag uptunnel in the plume. The percentage of moths doing so corresponded to the difference between the peak concentration within the plume and the background concentration of pheromone permeating the airstream. Moths also locked onto, and flew upwind along the pheromone-clean-air boundary formed along a pheromone-permeated side corridor. Because a similar response was observed along a horizontal edge between a pheromone-permeated floor corridor and clean air, we conclude that the intermittent stimulation at the edge perpetuated the narrow zigzagging response to pheromone.     

Interneurons sensitive to female pheromone in the deutocerebrum of the male silkworm moth, Bombyx mori

ABSTRACT. In response to minute quantities of female sex pheromone, the male silkworm moth, Bombyx mori L., walks upwind to locate the odour source. The axons of antennal receptors specific for the two known components of the pheromone terminate in the deutocerebrum. In this study, single interneurons were recorded extracellularly in the deutocerebrum of the male silkworm moth. Responses were characterized as the antennae were presented with puffs of clean air, or air containing either or both components of the female pheromone, bombykol and bombykal. An apparatus is described which added bombykol or bombykal to a constant air stream flowing over the antenna. Most units (87%) showed qualitatively different responses to bombykol and bombykal. A majority of the pheromone-sensitive units (65%) also showed mechanosensory responses to air puffs. Two units were recorded which were slightly inhibited by either bombykol or bombykal alone, but were excited by a mixture of the two.     

Sex pheromone and visual trap interactions in mate location strategies and aggregation by host-alternating aphids in the field

Abstract. Field observations were made on the responses of males and gynoparae of three host-alternating aphid species, the blackberry-cereal aphid, Sitobion fragariae (Walker), the bird cherry-oat aphid, Rhopalosiphum padi (L.) and the damson-hop aphid, Phorodon humuli (Schrank) to species-specific sex pheromones released from transparent and coloured water traps.Pheromone traps caught significantly more males than did control traps without pheromone, whereas transparent, light green, yellow and orange traps caught most insects.Measurements of the distance over which sex pheromones function indicated that male P.humuli detect the pheromone 2–6 m from the source and can fly upwind to a source in wind speeds of 0.7 m s^-1.In all three species significantly more gynoparae were caught in pheromone traps than in control traps, suggesting that pheromone released by adult sexual females may assist late-flying gynoparae to locate a suitable host plant on which to deposit their progeny.The response is relatively stronger for males than gynoparae, but the pheromones appear to act as both sex and aggregation pheromones.     

Pheromone-mediated optomotor anemotaxis and altitude control exhibited by male oriental fruit moths in the field

Abstract. In the field over short grass, pheromone-stimulated oriental fruit moth males, Grapholita molesta (Busck), flying under high windspeeds tended to steer courses more into the wind and to increase their airspeeds compared with those flying in low windspeeds.Thus, optomotor anemotaxis enabled the males to steer relatively consistent upwind track angles and to maintain an upwind progress of between c. 50–100 cm/s despite variable wind velocities.Zigzagging flight tracks were observed at both 10 m and 3 m from the source, as were tracks with no apparent zigzags.Transitions from casting to upwind flight or vice-versa were observed.The durations of the intervals between reversals during both upwind zigzagging flight and casting were consistent with those observed in previous wind-tunnel experiments.The control of altitude was more precise during upwind zigzagging flight than during casting.In general, the side-to-side deviations in the tracks were greater than the up-and-down deviations, with both the side-to-side and vertical distances and their ratios being consistent with previous wind-tunnel studies of pheromone-mediated flight.One difference between the field and laboratory flight tracks was that males in the field exhibited much higher airspeeds than in the wind tunnel.Males occasionally were observed to progress downwind faster than the wind itself, and further analysis showed that they were steering a downwind course in pheromone-free air following exposure to pheromone, which is the first time this has been recorded in moths.We propose that such downwind flight may aid in the relocation of a pheromone plume that has been lost due to a wind-shift, by enabling the moth to catch up to the pheromone as it recedes straight downwind away from the source.     

A non-anemotactic mechanism used in pheromone source location by flying moths

ABSTRACT. Male oriental fruit moths, Grapholitha molesta (Busck) (Tortricidae), continue to zigzag along a pheromone plume to the source in zero wind, if they have started flight with wind on. If the pheromone source is removed and the plume is hence truncated, moths flying in zero wind out of the end of the plume into clean air increase the width of their reversals and the angles of the straight legs of the tracks so they are more directly across the former wind line. Such moths reach the source less often than do those flying along a continuous plume. The males continue to zigzag up a plume in zero wind, apparently by a combination of sequential sampling of concentration along their path and the performance of an internal, self-steered programme of track reversals (zigzags) whose frequency increases with concentration. Visual feedback may aid in the still-air performance of the zigzags. We propose that both the sequential sampling (longitudinal klinotaxis) and self-steered counter-turning programme also are used in wind as well; anemotaxis apparently polarizes the direction of the zigzags to result in upwind displacement, and the narrow zigzags caused by the higher concentration in the plume keep the male 'locked on' to the odour.