An improved set of electron-THFA cross sections refined through a neural network-based analysis of swarm data.

We review experimental and theoretical cross sections for electron transport in α-tetrahydrofurfuryl alcohol (THFA) and, in doing so, propose a plausible complete set. To assess the accuracy and self-consistency of our proposed set, we use the pulsed-Townsend technique to measure drift velocities, longitudinal diffusion coefficients, and effective Townsend first ionization coefficients for electron swarms in admixtures of THFA in argon, across a range of density-reduced electric fields from 1 to 450 Td. These measurements are then compared to simulated values derived from our proposed set using a multi-term solution of Boltzmann's equation. We observe discrepancies between the simulation and experiment, which we attempt to address by employing a neural network model that is trained to solve the inverse swarm problem of unfolding the cross sections underpinning our experimental swarm measurements. What results from our neural network-based analysis is a refined set of electron-THFA cross sections, which we confirm is of higher consistency with our swarm measurements than that which we initially proposed. We also use our database to calculate electron transport coefficients in pure THFA across a range of reduced electric fields from 0.001 to 10 000 Td.

Fitness trade-off in peach-potato aphids (Myzus persicae) between insecticide resistance and vulnerability to parasitoid attack at several spatial scales.

Insecticide-resistant clones of the peach-potato aphid, Myzus persicae (Sulzer), have previously been shown to have a reduced response to aphid alarm pheromone compared to susceptible ones. The resulting vulnerability of susceptible and resistant aphids to attack by the primary endoparasitoid, Diaeretiella rapae (McIntosh), was investigated across three spatial scales. These scales ranged from aphids confined on individual leaves exposed to single female parasitoids, to aphids on groups of whole plants exposed to several parasitoids. In all experiments, significantly fewer aphids from insecticide-susceptible clones became parasitised compared to insecticide-resistant aphids. Investigations of aphid movement showed at the largest spatial scale that more susceptible aphids than resistant aphids moved from their inoculation leaves to other leaves on the same plant after exposure to parasitoids. The findings imply that parasitoids, and possibly other natural enemies, can influence the evolution and dynamics of insecticide resistance through pleiotropic effects of resistance genes on important behavioural traits.

Characterization of the M918T sodium channel gene mutation associated with strong resistance to pyrethroid insecticides in the peach-potato aphid, Myzus persicae (Sulzer).

Recent advances in the characterisation of insect sodium channel gene sequences have identified a small number of point mutations within the channel protein that are implicated in conferring target-site resistance to pyrethroid insecticides (so-called knockdown resistance or kdr). The L1014F (leucine-to-phenylalanine) mutation located in the centre of segment 6 of the domain II region (IIS6) of the sodium channel (the so-called kdr trait) has been detected in the peach-potato aphid, Myzus persicae (Sulzer), and is considered to be the primary cause of pyrethroid resistance in this species. Here we report on the characterisation of a second mutation, M918T (methione-to-threonine), within the nearby IIS4-S5 intracellular linker (the so-called super-kdr trait) in a field clone also possessing L1014F, with both mutations present in heterozygous form. The resistance phenotype of M. persicae clones possessing various combinations of L1014F and M918T to a wide range of pyrethroids (both Type I and II) was assessed in leaf-dip bioassays and to lambda-cyhalothrin applied at up to ten times the recommended field rate as foliar sprays to aphids feeding on whole plants. Bioassay results demonstrated that presence of both mutations was associated with extreme resistance to all the pyrethroids tested relative to aphids lacking the mutations. Furthermore, this resistance well exceeded that shown by aphids that were homozygous for L1014F but lacking M918T. However, pre-treatment with piperonyl butoxide in the leaf-dip bioassays failed to suppress pyrethroid resistance in aphids carrying one or both of the mutations. The relevance of these findings for monitoring and managing pyrethroid resistance in M. persicae populations in the field is discussed.

Regulation of pheromone biosynthesis in the "Z strain" of the European corn borer, Ostrinia nubilalis.

The regulation of pheromone biosynthesis by the neuropeptide PBAN in the Z strain of the European corn borer, Ostrinia nubilalis, was investigated using labeled intermediates. Injection of radiolabeled acetate showed PBAN did not influence the de novo synthesis of saturated fatty acids in the gland. When deuterium-labeled myristic acid was topically applied to the gland, females injected with PBAN produced more labeled pheromone than did control females, indicating that PBAN controls one of the later steps of pheromone biosynthesis. Although more myristic acid was Delta11-desaturated in the gland in the presence of PBAN, this was counterbalanced by less Delta11-desaturation of palmitic acid, indicating that desaturase activity did not change overall. This change in flux of myristic acid through to pheromone was shown to be caused by increased reduction of fatty acid pheromone precursors occurring in the presence of PBAN.

Evidence for two-step regulation of pheromone biosynthesis by the pheromone biosynthesis-activating neuropeptide in the moth Heliothis virescens.

The control of pheromone biosynthesis by the neuropeptide PBAN was investigated in the moth Heliothis virescens. When decapitated females were injected with [2-(14)C] acetate, females co-injected with PBAN produced significantly greater quantities of radiolabeled fatty acids in their pheromone gland than females co-injected with saline. This indicates that PBAN controls an enzyme involved in the synthesis of fatty acids, probably acetyl CoA carboxylase. Decapitated females injected with PBAN showed a rapid increase in native pheromone, and a slower increase in the pheromone precursor, (Z)-11-hexadecenoate. Total native palmitate and stearate (both pheromone intermediates) showed a significant decrease after PBAN injection, before their titers were later restored to initial levels. In contrast, the acyl-CoA thioesters of these two saturated fatty acids increased during the period when their total titers decreased. When a mixture of labeled palmitic and heptadecanoic (an acid that cannot be converted to pheromone) acids was applied to the gland, PBAN-injected females produced greater quantities of labeled pheromone and precursor than did saline-injected ones. The two acids showed similar time-course patterns, with no difference in total titers of each of the respective acids between saline- and PBAN-injected females. When labeled heptadecanoic acid was applied to the gland alone, there was no difference in titers of either total heptadecanoate or of heptadecanoyl-CoA between PBAN- and saline-injected females, suggesting that PBAN does not directly control the storage or liberation of fatty acids in the gland, at least for this fatty acid. Overall, these data indicate that PBAN also controls a later step involved in pheromone biosynthesis, perhaps the reduction of acyl-CoA moieties. The control by PBAN of two enzymes, near the beginning and end of the pheromone biosynthetic process, would seem to allow for more efficient utilization of fatty acids and pheromone than control of only one enzyme.

The attack of the clones: tracking the movement of insecticide-resistant peach-potato aphids Myzus persicae (Hemiptera: Aphididae).

Myzus persicae (Sulzer) collected in Scotland were characterized for four microsatellite loci, intergenic spacer fingerprints and the resistance mechanisms modified acetylcholinesterase (MACE), overproduced carboxylesterase and knockdown resistance (kdr). Microsatellite polymorphisms were used to define a limited number of clones that were either fully susceptible to insecticides or possessed characteristic combinations of resistance mechanisms. Within these clones, intergenic spacer fingerprints could either be very consistent or variable, with the latter indicating ongoing evolution within lineages, most likely derived from the same zygote. Two clones (termed A and B) possessed all three resistance mechanisms and predominated at sites treated with insecticides. Their appearance on seed potatoes and oilseed rape in Scotland in 2001 coincided with extensive insecticide use and severe control failures. Clones C, I and J, with no or fewer resistance mechanisms, were found in samples from 1995 and were dominant at untreated sites in 2001. A comparison of Scottish collections with those from other UK and non-UK sites provides insight into the likely origins, distribution and dynamics of M. persicae clones in a region where asexual (anholocyclic) reproduction predominates, but is vulnerable to migration by novel genotypes from areas of Europe where sexual (holocyclic) reproduction occurs.

The fate of topically applied fatty acids in the sex pheromone gland of the moth Heliothis virescens.

Deuterium-labeled hexadecanoic acid (D4-16:COOH), a sex pheromone biosynthetic intermediate, and heptadecanoic acid (D3-17:COOH), an acid that cannot be converted to sex pheromone, were topically applied to the pheromone gland of female Heliothis virescens, and the fate of the label determined. Both acids were incorporated similarly into the glycerolipids, with by far the greatest amount found in the triacylglycerols (TGs), and relatively small amounts found in other neutral and polar classes. For D4-16:COOH, the labeled pheromone precursor, (Z)-11-hexadecenoate, was also found predominantly in the TGs but relatively (compared to labeled hexadecanoate) high amounts were also found in the phospholipids. Within the TGs, both acids, as well as the pheromone precursor, were found almost exclusively on the sn-3 position of the glycerol backbone. This demonstrates that the major fate, in the glycerolipids, of free fatty acids is addition to 1,2-diacylglycerols. A relatively large amount of the applied acid was also found in the gland in the form of the acyl-CoA thioester. In a 24-h time-course study, this form remained at a relatively high level for the duration of the assay, and decreased at a rate comparable to the titer of this acid in the TGs, suggesting that titers of fatty acids in the glycerolipids and acyl-CoA thioesters may be in equilibrium. A time-course assay with D4-16:COOH demonstrated that peak pheromone titer after application was reached before peak titers of both total hexadecanoate and hexadecanoyl-CoA. Combined with a dose-response experiment, which showed that labeled pheromone titer did not increase above an applied concentration of 20 mg/ml, these data suggest that the final step in pheromone biosynthesis, reduction of Z11-16:Acyl-CoA, may be inhibited by increased acyl-CoA titers in the gland. Overall, our data are consistent with the glycerolipids modulating acyl-CoA concentrations in the pheromone gland.

Lipid analysis of the sex pheromone gland of the moth Heliothis virescens.

The sex pheromone gland of female Heliothis virescens was analyzed for fatty acid and lipid content. Base methanolysis of the gland showed a large amount of methyl (Z)-11-hexadecenoate (Z11-16:Acyl), the fatty acyl analog of the major pheromone component, (Z)-11-hexadecenal, as well as a small amount of methyl (Z)-11-octadecenoate. Methyl esters of various common fatty acids were also observed. HPTLC analysis of the glandular lipids revealed large quantities of triacylglycerols (TGs), and lesser amounts of 1,2-diacylglycerols (1,2-DGs), 2-monoacylglycerols (2-MGs), phosphatidyl ethanolamines, and phosphatidyl cholines. The greatest amount of Z11-16:Acyl in these lipids was in the TGs, with lesser amounts in the two phospholipid classes and only trace amounts in the other neutral lipids. The glands of females at various ages and photoperiodic times were extracted, fractionated into neutral and polar fractions by silica SPE, and fatty acid titers in these fractions determined. All fatty acids, but notably Z11-16:Acyl, showed significant total and neutral lipid fraction peaks at mid scotophase for 2-day-old females; a less dramatic, but significant, Z11-16:Acyl peak in the polar fraction was also observed. However, only a relatively small proportion (<50%) of this acid was recovered from the silica at all times. This "non-recoverable" Z11-16:Acyl showed a dramatic and significant peak at mid scotophase for 2-day females, corresponding roughly with maximal pheromone titer. All other acids in the gland were recovered in high proportions, and their respective "non-recoverable" titers were not different at any of the times analyzed. Based on previous work, this non-recoverable Z11-16:Acyl is likely the CoA ester. Therefore, it appears that the pheromone gland of H. virescens maintains pools of Z11-16:Acyl in both CoA ester and TG forms, which are available for biosynthesis of pheromone. These pools are greatest during maximal pheromone production when the biosynthetic enzymes, possibly the fatty acid reductase, are unable to utilize rapidly enough the quantities of Z11-16:Acyl biosynthesized.

Identification of sex pheromone components of the painted apple moth: a tussock moth with a thermally labile pheromone component.

The sex pheromone of the painted apple moth, Teia anartoides (Lymantriidae) was investigated using GC-EAD and GC-MS analysis, derivatization, TLC analysis, and field cage and field trapping bioassays. The major sex pheromone components were identified as (6Z,9Z)-henicosa-6,9-dien-11-one and (6Z,9Z)-henicosa-6,9-diene. Other minor components of pheromone gland extracts included (6Z)-9R, 10S-epoxyeicos-6-ene, (6Z)-9R,10S-epoxyhenicos-6-ene, (6Z,9Z)-henicosa-6,9-dien-11-ol, (6Z)-henicos-6-en-11-one, and (6Z, 8E)-henicosa-6,8-dien-11-one, but the roles of these minor components remain equivocal. In field cage and field experiments, a blend of all seven identified components [(6Z,9Z)-henicosa-6,9-dien-11-one (relative amount 100), (6Z,9Z)-henicosa-6,9-diene (100), (6Z)-9R,10S-epoxyeicos-6-ene (5), (6Z)-9R,10S-epoxyhenicos-6-ene (10), (6Z,9Z)-henicosa-6,9-dien-11-ol (5), (6Z)-henicos-6-en-11-one (1), and (6Z,8E)-henicosa-6,8-dien-11-one (25)] was as attractive to males as calling females, but tests with blends of the major component(s) with subsets of the minor components did not produce consistent results that unequivocally showed the various minor components to be critical components of the active blend. (6Z,9Z)-henicosa-6,9-dien-11-one is thermally labile and rearranges to (6Z,8E)-henicosa-6,8-dien-1-one and other products at ambient temperature, rendering the synthetic pheromone lure inactive after two days of field exposure.

Reduced response of insecticide-resistant aphids and attraction of parasitoids to aphid alarm pheromone; a potential fitness trade-off.

Response to the alarm pheromone, (E)-beta-farnesene, produced by many species of aphids, was assessed in laboratory bioassays using an aphid pest, Myzus persicae (Sulzer), and its primary endoparasitoid, Diaeretiella rapae (McIntosh). This was done in three separate studies, the first investigating responses of a large number of M. persicae clones carrying different combinations of metabolic (carboxylesterase) and target site (kdr) insecticide resistance mechanisms, and the other two investigating the responses of young virgin female adult parasitoids. In M. persicae, both insecticide resistance mechanisms were associated with reduced repellence suggesting that each has a pleiotropic effect on aphid behaviour. In contrast, D. rapae females were attracted to the alarm pheromone source. The implications of this apparent fitness trade-off for the evolution and dynamics of insecticide resistance, and the potential for using beneficial insects to combat resistance development are discussed.

Fatty acid and sex pheromone changes and the role of glandular lipids in the Z-strain of the European corn borer, Ostrinia nubilalis (Hübner).

Lipids in the sex pheromone gland of females of the Z-strain of Ostrinia nubilalis were analyzed for fatty acyl pheromone analogs (FAPAs) and other potential biosynthetic intermediates. More than 80% of the FAPAs were found in the triacylglycerols (TGs), with smaller amounts found in the phosphatidyl cholines, ethanolamines, and serines. Analysis of the TGs by lipase revealed that the two FAPAs were distributed fairly evenly among all three stereospecific positions. Comparison of changes in titers of key glandular fatty acids with those of pheromone components, with respect to photoperiodic time and age of females, showed that both FAPA and pheromone titers exhibited a cyclical pattern with peaks in the scotophase and valleys in the photophase. However, whereas pheromone titer tended to peak in the first half of the scotophase, FAPA titer peaked at the end of the scotophase. Significantly, the titer of the FAPA of the minor component, (E)-11-tetradecenyl acetate (3% of pheromone), was always much greater than the titer of the FAPA of the major component, (Z)-11-tetradecenyl acetate (97%), of the pheromone. Titer of myristate, an intermediate in pheromone biosynthesis, was also higher during the scotophase than the photophase. However, myristate titer showed a pronounced dip in the middle of the scotophase. These data suggest two roles for glandular lipids in sex pheromone biosynthesis in O. nubilalis. Firstly, they remove excess FAPA of the minor component so the fatty acid reductase system is not presented with a high ratio of this isomer (which would otherwise result from the reductase's own selectivity), which could cause changes in the final pheromone ratio. Secondly, hydrolysis of the large amounts of stored saturated fatty acids from the TGs may provide substrate for pheromone biosynthesis.

The effects of topical application of various fatty acids on pheromone and glandular lipid biosynthesis in the moth Heliothis virescens.

Binary mixtures of deuterium-labeled palmitic acid and an excess of different fatty acids were applied to the sex pheromone gland of female Heliothis virescens and the effects on the terminal steps of pheromone biosynthesis, including incorporation of fatty acids into the glandular lipids, observed. Relative to labeled palmitic acid applied alone, application of all the binary mixtures resulted in decreased levels of the labeled pheromone component, (Z)-11-hexadecenyl acetate (Z11-16:OAc), but there was generally no decrease in the amounts of labeled pheromone precursor, (Z)-11-hexadecenoate, nor labeled palmitate in the glandular lipids. These data suggest that the excess of fatty acid in the gland inhibits Delta11-desaturation. However, in the case of excess myristoleic acid, the amount of labeled (Z)-11-hexadecenoate increased significantly, suggesting that this acid inhibited fatty acid reduction. Dose-response tests with certain of the fatty acids were consistent with the above interpretations and further indicated that the gland had a high capacity for rapidly activating and incorporating excess fatty acids into the glandular lipids. Finally, application of the various fatty acids resulted in increased levels of these acids in the gland and, in the cases of myristoleic, palmitoleic and myristic acids, it also resulted in increased levels of the corresponding aldehydes, which had previously been detected in the gland of female H. virescens. This suggests that the fatty acid reductase in H. virescens is not highly specific for the major component, and that the final ratio of pheromone components is determined in part by the availability of their corresponding fatty acids in the gland.

Analogous pleiotropic effects of insecticide resistance genotypes in peach-potato aphids and houseflies.

We show that single-point mutations conferring target-site resistance (kdr) to pyrethroids and DDT in aphids and houseflies, and gene amplification conferring metabolic resistance (carboxylesterase) to organophosphates and carbamates in aphids, can have deleterious pleiotropic effects on fitness. Behavioural studies on peach-potato aphids showed that a reduced response to alarm pheromone was associated with both gene amplification and the kdr target-site mutation. In this species, gene amplification was also associated with a decreased propensity to move from senescing leaves to fresh leaves at low temperature. Housefly genotypes possessing the identical kdr mutation were also shown to exhibit behavioural differences in comparison with susceptible insects. In this species, resistant individuals showed no positional preference along a temperature gradient while susceptible genotypes exhibited a strong preference for warmer temperatures.

Fatty acyl pheromone analogue-containing lipids and their roles in sex pheromone biosynthesis in the lightbrown apple moth, Epipyhas postvittana (Walker).

The pheromone gland of the moth Epiphyas postvittana was analysed for lipids containing the fatty acyl pheromone analogue (FAPA) of the component, (E)-11-tetradecenyl acetate. The FAPA was found predominantly in the triglycerides (TGs), and to a lesser extent in the choline phosphatides. The FAPA was found to be exclusively on the sn-1 or sn-3 position (probably the latter) of the TGs. When pheromone gland lipid extracts were eluted through silica solid phase extraction, a significant proportion of the FAPA was not recovered. Changes in titre of this non-recoverable FAPA paralleled changes in pheromone titre in females. In contrast, changes in recoverable FAPA (mostly in the TGs) titre showed a gradual increase with time after eclosion. The properties of this non-recoverable FAPA were consistent with it being the CoA ester of the FAPA. Thus, it appears that the FAPA-CoA ester is the immediate lipid precursor of the pheromone, and that the FAPA-containing TGs are formed by reaction of the FAPA-CoA with 1,2-DGs, as a consequence of the rate-limiting reduction of the FAPA-CoA. Finally, injection of PBAN into females decapitated for 3 days resulted in a decrease in recoverable FAPA and an increase in non-recoverable FAPA, suggesting that PBAN influences the lipolysis of TGs. Overall these data suggest that there are two routes for biosynthesis of the pheromone component E11-14:OAc in E. postvittana: a de novo route, directly via the CoA esters of the various fatty acid intermediates, and a less direct route via the lipolysis of FAPA-containing TGs.

Periodicity of sex pheromone biosynthesis, release and degradation in the lightbrown apple moth, Epiphyas postvittana (Walker).

Pheromone titer in moths is a product of three processes occurring in or at the surface of the pheromone gland: biosynthesis, release, and intraglandular degradation, of pheromone. Changes in titers of sex pheromone, the fatty acyl pheromone analog (FAPA), and tetradecanoate, a pheromone biosynthetic intermediate, were studied in detail in the lightbrown apple moth, Epiphyas postvittana (Walker). Although changes in the pheromone titers in a day were relatively small, with the peak titer being 2-3 times greater than that at the trough, pheromone titer did show a distinct diel periodicity. Titer of the FAPA showed a similar, but less variable, diel pattern, but tetradecanoate titer showed little or no diel pattern. The pattern of pheromone titer suggested that females biosynthesize pheromone at two different rates during the photoperiod: a high rate during the latter half of the photophase and most of the scotophase, which is associated with a high pheromone titer, and a low rate throughout the first half of the photophase, which is associated with a low titer. Consistent with data on commencement of copulation, pheromone was released from the second hour of the scotophase through to the eighth hour. Pheromone release rate during this period appeared to be similar to the rate of pheromone biosynthesis. In contrast to the other two processes, pheromone degradation did not appear to have a diel pattern. Females decapitated at different times of the photoperiod showed a similar decline in pheromone titer, consistent with the reaction kinetics being first order in pheromone titer.

A sodium channel point mutation is associated with resistance to DDT and pyrethroid insecticides in the peach-potato aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae).

The voltage-gated sodium channel is the primary target site of DDT and pyrethroid insecticides, and point mutations in the domain II region of the channel protein have been implicated in the knockdown resistant (kdr ) phenotype of several insect species. Here, we report that one of these mutations, a leucine-to-phenylalanine replacement in transmembrane segment IIS6, is also found in certain insecticide-resistant clones of the peach-potato aphid, Myzus persicae. The mutation was present in four clones with amplified E4 esterase genes, but was absent from both susceptible clones and those with amplified FE4 genes. The inferred presence of kdr-type resistance in the four E4 clones was subsequently confirmed by bioassays that showed this to be the primary mechanism of resistance to deltamethrin and DDT, although the esterase-based mechanism also contributes to the overall level of deltamethrin resistance. The kdr mutation on its own conferred 35-fold resistance to deltamethrin and this was enhanced up to 540-fold when it was present in a high (E4) esterase background. The esterase (FE4) mechanism was far less effective without the kdr mutation, conferring just 3-4-fold resistance to deltamethrin. These findings, and the linkage disequilibrium of the kdr mutation within clones overproducing the E4 esterase, have important implications for the evolution of resistance in this insect and for the use of pyrethroid sprays in the management of M. persicae populations in the field.

Behavioral manipulation methods for insect pest-management.

We discuss methods using stimuli to manipulate behavior of a pest for the purpose of protecting a valued resource. The methods are divided into two categories: those that manipulate behavior over a long distance, e.g. volatile chemicals, visual, and auditory stimuli, and those that manipulate behavior at a short distance (<1 cm), e.g. involatile chemicals. Particular emphasis is placed on methods that have been developed through studies of pest behavior and on combining stimuli to increase efficacy. Future prospects for behavioral manipulation methods in pest management are discussed.

Change in reductase activity is responsible for senescent decline in sex pheromone titre in the lightbrown apple moth, Epiphyas postvittana (Walker).

Sex pheromone titre in the tortricid moth Epiphyas postvittana follows a pattern commonly observed in other species of moths: an increase to a peak some time after eclosion (2-3days), and then a slow decline as the female ages. Previous work has shown that this decline is not regulated by the pheromone biosynthesis activating neuropeptide PBAN. Using in vivo and in vitro enzyme assays, and fatty acid methyl ester (FAME) analyses of pheromone precursors in the gland, we have investigated this senescent decline in pheromone titre. The enzyme assays have shown that in older females the fatty acid reductase and fatty acid synthesis enzyme systems decrease in activity (relative to younger females), whereas other enzyme systems involved in pheromone biosynthesis, including limited beta-oxidation (2-carbon chain-shortening), (E)-11-desaturation, and acetylation (by an acetyl transferase) remain unchanged in their activity. Of the two enzymatic processes involved, the more important one contributing to the decline appears to be the fatty acid reductase. This is consistent with FAME analyses of pheromone glands in old and young females, which show little difference in levels of saturated FAME, but a significant increase in the level of the putative precursor, (E)-11-tetradecenoate, of the sex pheromone component (E)-11-tetradecenyl acetate. Thus, this decline in fatty acid reductase activity results in a buildup of the precursor as the female ages. The near ubiquity of fatty acid reductases in moth sex pheromone systems suggests that this may be a common mechanism for the senescent decline of sex pheromone titre in moths.

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.

Effect of design of a sex-pheromone-baited delta trap on behavior and catch of maleEpiphyas postvittana (Walker).

Through field trials and wind-tunnel studies, we have demonstrated that certain structural features of a sex-pheromone-baited delta trap affect catch of light-brown apple moth,Epiphyas postvittana, males, by influencing behaviors used to enter and exit the trap. Field catch of males was dependent upon length (and width) of the trap, with increases in length yielding linear increases in catch. In the wind tunnel, similar numbers of males entered the two traps, but significantly fewer males exited the longer trap within 1 min after entering it. Although males landed on the sticky surface at similar distances from the downwind entrances of the traps, they were stuck farther upwind on the longer trap. Thus, it is probable that the increase in field catch with increase in trap length relates to the increase in distance (and hence time) that males walk on the sticky surface, towards the pheromone source, before attempting to exit. The bottom barriers (as well as additional barriers at the top and sides) at the entrances of the trap also significantly influenced trap catch. The barriers apparently influence trap catch in two ways. Firstly, they hinder the exit of males from the trap, thus diverting males back into the trap and increasing their chance of being caught. Secondly, they influence where the male lands on the sticky surface; with higher barriers, males land farther upwind (i.e., nearer the source), and thus farther from an exit. Finally, as the source was suspended higher above a horizontal surface, greater numbers of males landed on the source. This result shows that the position where a maleE. postvittana lands is influenced by the relationship of the source to the surface and suggests that trap catch of males may similarly be influenced (i.e., by inducing males to land farther from the exits).