Mechanisms of Pheromone Communication Disruption in Choristoneura rosaceana Exposed to Microencapsulated (Z)-11-tetradecenyl Acetate Formulated with and Without Horticultural Oil

Flight tunnel and electrophysiological assays with male Choristoneura rosaceana Harris (Lepidoptera: Tortricidae) were conducted to investigate the non-competitive mechanisms of communication disruption caused by microencapsulated (MEC) (Z)-11-tetradecenyl acetate (3M MEC-LR) formulated with and without horticultural oil (Purespray Green). Male C. rosaceana were exposed for 1 h by resting on metal surfaces sprayed with either water, 2% oil in water, MEC-LR in water, or MEC-LR + 2% oil in water. In one experiment, sprayed surfaces were allowed to age up to 47 days in a laboratory fume hood prior to moth exposure to examine the effect of ageing on the disruptive efficacy of the formulations. In flight tunnel assays with calling females, males exposed to MEC or MEC + oil treatments for 1 h were significantly disrupted up to 1 h after exposure, and both treatments were effective for 47 days. Electroantennograms revealed no reduction in antennal sensitivity when measured 75 s after a 1-h exposure. Collectively, these results support habituation as a key mechanism of communication disruption for C. rosaceana exposed to the MEC formulations tested here. Male proximity to the pheromone-treated surfaces appears to be important in maintaining a disruptive effect as MEC treatments age and pheromone release rates decline. The addition of 2% oil increased the number of microcapsules deposited on sprayed surfaces and caused a moderate but significant increase in the disruptive effect of the MEC formulation. A better understanding of the mechanisms that underlie disruption by MEC formulations, and how an adjuvant like horticultural oil may enhance these mechanisms, may lead to improvements in this technology.     

Sex pheromone components of the obliquebanded leafroller moth,Choristoneura rosaceana

Previously, (Z)-11-tetradecen-1-yl acetate was reported as the sex pheromone of the obliquebanded leafroller,Choristoneura rosaceana (Harris). A mixture of 92% (Z)-11-tetradecen-1-yl acetate and 8% (E)-tetradecen-1-yl acetate has been reported as an attractant for this insect. (E)-11-tetradecen-1-yl acetate and (Z)-11-tetradecen-1-ol have now been isolated and identified from female tip extracts and shown to be behaviorally active; in addition, there is some evidence for the presence of (E)-11-tetradecen-1-ol in these extracts. TheZE ratio for both the acetates and the alcohols in these tip extracts was about 982; the proportion of alcohols was rather variable, but the acetates always predominated. Maximum trap catches fn the field during the course of this study were obtained with 5 mg of 955 (Z)(E)-11-tetradecen-1-yl acetates containing 0.5–10% (Z)-11-tetradecen-1-ol (approx. 1%Eisomer) in polyethylene caps. No evidence was found for the presence of (Z)-11-tetradecenal in female tip extracts, and this compound was found to be ineffective in increasing trap catches.Lepidoptera: Tortricidae: Tortricinae.This research was supported in part by the Rockefeller Foundation, and by NSF Grant No. GB-38020.     

Sex pheromone components of the oblique-banded leafroller,Choristoneura rosaceana in the Okanagan Valley of British Columbia

(Z)-11-Tetradecen-1-yl acetate, (E)-11-tetradecen-1-yl acetate, and (Z)-11-tetradecen-1-ol were previously reported as the sex pheromone in New York strains of the oblique-banded leafroller,Choristoneura rosaceana (Harris), and (E)-11-tetradecen-1-ol was tentatively identified in female tip extracts. For Okanagan Valley strains ofC. rosaceana, an additional component, (Z)-11-tetradecenal, was identified from female tip extracts by split-less capillary gas-liquid chromatography and mass spectroscopy and was strongly stimulatory in electroantennogram studies. In field tests, 3 mg of 96.521.5 (Z)-11-tetradecen-1-yl acetate, (E)-11-tetradecen-1-yl acetate, and (Z)-11-tetradecen-1-ol (containing approx. 1%E isomer) was not as attractive as female-baited traps, and significant numbers of European leafrollerArchips rosanus L. were attracted. The above blend with 1% (Z)-11-tetradecenal added was significantly more attractive than traps baited with femaleC. rosaceana, butA. rosanus males were still attracted. Increased percentages of (Z)-11-tetradecenal up to 4% caused increased catches ofC. rosaceana and decreased catches ofA. rosanus. Low amounts of (E)-11-tetradecen-1-ol and (E)-11-tetradecenal male also contribute to increased specificity of the synthetic pheromone blend toC. rosaceana.Lepidoptera: Tortricidae.Contribution No. 638, Agriculture Canada, Research Station, Summerland, British Columbia, Canada.     

Spruce budworm (Choristoneura fumiferana) pheromone chemistry and behavioral responses to pheromone components and analogs

This paper reviews the sex pheromone chemistry and pheromone-mediated behavior of the spruce budworm and related coniferophagous (Choristoneura) budworms. InC. fumiferana, temporal changes in pheromone-gland monounsaturated fatty acids (pheromone precursors) enable the prediction of the primary sex pheromone components. This technique may also be applicable for predicting additional pheromone components. Tetradecanal (14 Ald), previously shown to enhance close-range precopulatory behavior, lowers the threshold of response by males for upwind flight to a pheromone-component source. Spruce budworm males maintain upwind flight to 955 (E/Z)-1,12-pentadecadiene (diolefin analog) after initiating upwind flight to a primary-component pheromone source (955E/Z11-14Ald). This is the first demonstration of apparently normal male flight responses to a pheromone analog.Lepidoptera: Tortricidae     

Disruption of male spruce budworm orientation to calling females in a wind tunnel by synthetic pheromone

Male spruce budworm [Chorisloneura fumiferana (Clem.)] moths were held for 3 hr in a wind tunnel and subjected to various concentrations of background synthetic pheromone. They were then exposed to calling females and their response was recorded. The background pheromone was presented either as discrete turbulent plumes or as a uniform permeation throughout the tunnel. The numbers of males wing-fanning and flying in response to the calling females decreased as the concentration of background pheromone increased. Of the males which flew, a higher proportion progressed upwind in the discrete plumes than in the uniform permeation, an indication that structure in the pheromone cloud is necessary for upwind progression. In both discrete plumes and uniform permeation fewer males were able to locate the females (i.e., disruption was greater) as the concentration of synthetic pheromone increased, but for the same total release rates, disruption was greater when the synthetic pheromone was released in discrete plumes rather than in a uniform permeation. This implies that disruption which involves luring males to sources of synthetic pheromone is more effective than masking female plumes by uniform permeation and suggests that it is more efficient to release pheromone from a few potent sources than from numerous low-potency sources.     

Effects of Tansy Essential Oil from Tanacetum vulgare on Biology of Oblique-Banded Leafroller, Choristoneura rosaceana

The formulations TSW (tansy essential oil, surfactant, distilled water) and TSWD (tansy essential oil, surfactant, distilled water, dillapiol) increased the feeding rate of oblique-banded leafroller (OBLR) larvae with increasing concentrations of tansy essential oil. When tansy essential oil was continuously present at 0.1% and 1% in the diet, all OBLR larvae from susceptible and resistant populations died. The presence of 0.01% tansy essential oil in the diet affected female pupal weight, but not larval weight gain, larval developmental time and male pupal weight. Our results suggest that the concentration of tansy essential oil that significantly affects the developmental variables should be between 0.01% and 0.1% of tansy essential oil. Residues of the formulation TE (tansy essential oil, ethanol 95%) deterred oviposition of OBLR female.     

Changes in pheromone titer of oblique-banded leafroller,Choristoneura rosaceana,virgin females as a function of time of day,age, and temperature

Under a 16:8 hr light-dark photoperiod and 20C constant temperature, the titers of (Z)-11-tetradecenyl acetate (Z11–14:Ac), (E)-11-tetradecenyl acetate (E11–14:Ac) and (Z)-11-tetradecenol (Z11–14:OH) produced by different-agedChoristoneura rosaceana virgin females varied significantly during the scotophase, with the maximum titer occurring before the onset of calling in day-0 and day-3 females, while in day-5 females the titer remained constant throughout the calling period. There was a significant decrease in the titer of all pheromone components with age, explaining the lesser attractiveness of day-5 females relative to day-0 and day-3 females observed in the field. Under a cold thermocycle simulating condition during the second flight period in the fall, the titers of all pheromone components did not vary with time of day. There was a significant decrease in the amount ofZ11–14:Ac with age but no changes occurred in the minor components. Furthermore, for any given age tested, the amount of each component produced during the period of maximal calling activity remained relatively similar at the two temperature regimes. However, as with the expression of calling behavior, pheromone production was initiated earlier at cooler than at warmer temperatures. At both temperature regimes, female age and time of day influenced the ratio of each pheromone component. These results are discussed in relation to the hypothesis that by calling earlier, less attractive older females may increase their probability of mating.     

The Effect of Atmospheric Pheromone Concentrations on Behavior of Lightbrown Apple Moth in an Apple Orchard

A previously validated Lagrangian model was used to estimate the threshold of atmospheric pheromone concentration required to prevent trap catch and wing fanning in mating disruption plots in an apple orchard. Electroantennogram (EAG) traces of 10 min duration were recorded, along with supporting meteorological data needed for the model, to better define the conditions in which successful mating disruption will occur. Pheromone was released from polyethylene tubing dispensers into orchard blocks treated with 10, 100, 1000, and 2000 dispensers/ha. Predicted dusk concentrations of atmospheric pheromone at a height of 1.85 m varied nightly between 4 and 90 ng pheromone/m³ (in plots treated with 1000 dispensers/ha) over 11 weeks. Disruption of traps baited with 1000-g pheromone lures followed an asymptotic curve with predicted concentration, but they did not show a significant effect of trap height (1.5 and 3.0 m). Wing fanning was reduced by increasing the density of dispensers, but was not completely eliminated even at 1000 dispensers/ha. At this density, the concentrations were usually <16 ng pheromone/m³. Electroantennogram recordings of 10 min duration showed a higher frequency of pheromone pulses in plots treated with more point sources per hectare. There was also a positive correlation between the number of pulses recorded by the EAG and predicted concentration for plots treated with 1000 or 2000 dispensers/ha. These results give added support to our model of pheromone release and transport in treated apple orchards.     

Sex pheromone of a conifer-feeding budworm,Choristoneura retiniana,Walsingham

Sex pheromone components collected from femaleChristoneura retiniana included 11-tetradecenyl acetates and alcohols. The major component wasE-11-tetradecenyl acetate (E11–14Ac) with a lesser amount ofZ isomer necessary to induce male response. A 928 ratio ofE,Z11–14 Ac appeared optimal. The alcohol component was present at about 10% of the total pheromone mixture, and traps baited with acetates plus alcohol surpassed unmated females in their degree of attractiveness. Chemical analysis indicated a 9:1 ratio of theE — Z isomers of 11-tetradecenyl alcohol (11–14ol) pheromone components, although bait formulations containing a predominance of either theE orZ isomers were equally successful in field bioassays. Based on male response to traps,E- andZ11–14Ac (928E — Z) are essential pheromone components for long-range sex attraction. The 11–14 ol enhanced attraction when added at 10% of the total pheromone blend.This paper reports the results of research only. Mention of a commercial product does not constitute a recommendation by the U.S. Department of Agriculture.     

Attraction of male spruce budworm moths,Choristoneura fumiferana (Clemens), to pheromone-baited traps in small-tree thinnings

Mean catches of spruce budworm,Choristoneura fumiferana (Clemens), moths were not significantly different among four small-tree thinning treatments of young spruce-fir-hemlock regeneration. Significant inverse relationships were found between trap catches and distances to nearby spruce-fir-hemlock overstory. Prevailing wind directions indicated that moths were attracted anemotactically to upwind pheromone sources. No definite trends were detected between catches and temperature or precipitation.Lepidoptera: Tortricidae.A contribution to the Canada/ United States (CANUSA) Spruce Budworms Program. Mention of proprietary products does not constitute an endorsement of those products by the USDA.     

Identification of sex pheromone components of jack pine budworm,Choristoneura pinus pinus freeman

Chemical identification and field-trapping experiments have shown that a blend of 8515 (E,Z)-11-tetradecenyl acetates and 8515 (E,Z)-11-tetradecen-1-ols (in a 91 ratio) are female sex pheromone components for jack pine budworm,Choristoneura pinus pinus. This blend of chemicals, formulated in PVC (0.1 %, w/w) sources is as effective a trap bait as virgin females. Preliminary wind tunnel observations have indicated that this blend, effective as a trap bait, is not equivalent to females.Lepidoptera: Tortricidae.     

Responses of Tea Tussock Moth, Euproctis pseudoconspersa, to Its Pheromone, (R)-10,14-Dimethylpentadecyl Isobutyrate, and to the S-Enantiomer of Its Pheromone

Field trials were conducted with each synthetic enantiomer (>98% ee) and blends of the two synthetic enantiomers of the female-produced sex pheromone (10,14-dimethylpentadecyl isobutyrate) of the tea tussock moth, Euproctis pseudoconspersa. Male moths were attracted to each enantiomer alone and to various blends of them. Short syntheses of both enantiomers of the pheromone from commercially available (R)- and (S)-citronellyl bromide and a method of checking the enantiomeric purity of the citronellyl bromide enantiomers are described.     

Chemistry and field evaluation of the sex pheromone of western spruce budworm,Choristoneura occidentalis,Freeman

Chemical analyses and field bioassays showed a mixture of 92% (E)- and 8% (Z)-11-tetradecenal to be the sex attractant pheromone of the western spruce budworm,Choristoneura occidentalis Freeman. Females were also found to emit small amounts of the corresponding acetates and alcohols, but these components were not active in the bioassay. In contrast, in whole female tip extracts, (E)-11-tetradecenyl acetate predominates, and aldehyde pheromone components are present in lesser quantities, suggesting that final biosynthesis of pheromone takes place just prior to or during emission. At release rates approximating that of the female (2–4 ng/hr), a 928EZ blend of the synthetic aldehydes was at least as attractive as live females. Addition of the corresponding acetates or alcohols up to 50% of the aldehyde content did not significantly enhance or inhibit attraction. No major differences were apparent in pheromone production of females from a laboratory stock or from field collections from diverse geographic locations ranging from Colorado to British Columbia.     

Sex Pheromone of the Peach Aphid, Tuberocephalus momonis, and Optimal Blends for Trapping Males and Females in the Field

Chemical analysis of the volatiles released by sexual females (oviparae) of the peach aphid, Tuberocephalus momonis, identified two ubiquitous aphid sex pheromone components, (4aS,7S,7aR)-nepetalactone and (4aS,7S,7aR)-nepetalactol, in a ratio of 4 : 1. In field trials in Korea, employing traps releasing the two compounds in differing ratios, the most effective sex pheromone blend for trapping male T. momonis was found to be 85 : 15 nepetalactone–nepetalactol. Surprisingly, large numbers of presexual females (gynoparae) of this species were also collected when the catching rates were highest. In addition to T. momonis, over 20 other species of aphids were caught, particularly Myzus lythri, M. dycei, Lachnus tropicalis and M. persicae, in descending order of abundance.     

Seasonal Variation of Pheromone Concentration in Mating Disruption Trials Against European Grape Vine Moth Lobesia botrana (Lepidoptera: Tortricidae) Measured by EAG

Spatial and temporal distributions and relative concentrations of the pheromone of the European grape vine moth Lobesia botrana (Lepidoptera: Tortricidae) were measured in mating disruption trials with electroantennograms. Measurements were carried out over several years during the flight season of the two generations of this pest insect. In three consecutive years significantly higher mean relative pheromone concentrations were measured in summer during the flight of the second generation of L. botrana than in spring during the flight of the first generation (P < 0.001). The relative pheromone concentrations in 1989 reached a maximum between late July and early August, when the highest mean daily temperatures were registered. In addition to the differences in mean relative pheromone concentrations, the spatial and temporal distributions of the pheromone differed significantly between spring and summer. EAG recordings taken in summer showed high, uniform pheromone concentrations in the treated plots, whereas in spring strong temporal and spatial fluctuations were recorded. In a vineyard defoliated by a hailstorm, the mean relative pheromone concentrations measured in summer were not significantly different from those measured in spring (P > 0.05), but were significantly lower than those of a nearby intact vineyard (P < 0.001). The results provided additional evidence that foliage is an important parameter determining mean pheromone concentrations and temporal and spatial distribution of pheromone in mating disruption trials.     

Variables Affecting Pheromone Concentration in Vineyards Treated for Mating Disruption of Grape Vine Moth Lobesia botrana

Airborne pheromone concentration in a field is one of the most important variables for the successful application of mating disruption in pest control. In the present paper, we estimated the pheromone concentration with field EAG recordings in vineyards and investigated parameters affecting concentration. Pheromone concentration showed a positive correlation with number of dispensers per hectare (= number of point sources). A twofold increase in the absolute number of dispensers per hectare with a constant number of point sources (two dispensers at the same location) did not significantly affect relative pheromone concentration. Measurements carried out in plots where dispensers had been applied at different heights showed highest relative pheromone concentrations in plots with dispensers at 0.1 m and 1.4 m above the ground. Those concentrations were not significantly different from each other, but were significantly higher than in plots where dispensers had been placed at a height of 2 m. Foliage of grape vines substantially affected the development of high pheromone concentrations. In summer, in vineyards with fully developed leaf canopy, significantly higher pheromone concentrations could be measured shortly after application of the dispensers compared to vineyards in spring with sparse vegetation. The decline of pheromone concentrations after removal of dispensers is significantly prolonged in full vegetation, showing the impact of plant canopy on pheromone concentrations. In contrast, ground cover between the grapevine rows did not significantly affect mean pheromone concentrations.     

Metabolism of pheromone components and analogs by cuticular enzymes ofChoristoneura fumiferana

In vivo incubation ofChoristoneura fumiferana with (E)-11-tetradecenal, the major pheromone component, resulted in adsorption followed by transformation to (E)-11-tetradecenoic acid, the only observed metabolite. Antennae, legs, and wings from both sexes were investigated for enzyme activity. Specificity studies with the aldehyde dehydrogenase system showed that the enzyme prefers aldehydes of 14-carbon chain length or shorter.     

Pheromone Communication in the Honeybee (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.)

Recent studies have demonstrated a remarkable and unexpected complexity in social insect pheromone communication, particularly for honeybees (Apis mellifera L.). The intricate interactions characteristic of social insects demand a complex language, based on specialized chemical signals that provide a syntax that is deeper in complexity and richer in nuance than previously imagined. Here, we discuss this rapidly evolving field for honeybees, the only social insect for which any primer pheromones have been identified. Novel research has demonstrated the importance of complexity, synergy, context, and dose, mediated through spatial and temporal pheromone distribution, and has revealed an unprecedented wealth of identified semiochemicals and functions. These new results demand fresh terminology, and we propose adding “colony pheromone” and “passenger pheromone” to the current terms sociochemical, releaser, and primer pheromone to better encompass our growing understanding of chemical communication in social insects.     

Effect of foliage proximity on attraction ofChoristoneura occidentalis andC. retiniana (Lepidoptera: Tortricidae) to pheromone sources

Pheromone-baited traps located close to both host and nonhost crowns were more attractive than traps located between crowns for bothC. occidentalis Freeman andC. retiniana (Walsingham) at both 10 m and at 1.5 m above the ground. At 10 m height, traps located in host foliage were more attractive than those located in nonhost foliage, but at 1.5m height there was no significant difference. These results were obtained for both dense and sparse populations ofC. occidentalis and sparse populations ofC. retiniana. We conclude that the tree species on which a virgin female is located is not an important factor restricting mating between closely relatedChoristoneura spp. Also, the tree species on which a trap is located may not be an important factor that must be standardized in developing pheromone monitoring systems forC. occidentalis andC. retiniana.     

Behavioral effects of secondary components of sex pheromone of western spruce budworm (Choristoneura occidentalis) free

In a laboratory wind tunnel, upwind flight and close-range orientation to a pheromone source by maleC. occidentalis were facilitated by the addition of the secondary componentsE/Z11-tetradecen-1-ol acetate (8911) andE/Z11-tetradecen-1-ol (8515) to low source concentrations of the primary components,E/Z11-tetradecenal (928). Male responses to the blends tested never equalled their responses to virgin females. The primary components alone, when released at a rate similar to that of a calling female, never elicited male upwind flight or source location. However, the addition of the secondary components enhanced these behavioral sequences.Lepidoptera: Tortricidae.