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.     

Effect of a Rugulosin-producing Endophyte in Picea glauca on Choristoneura fumiferana

Disease-free second instar Choristoneura fumiferana (eastern spruce budworm) were placed on trees infected with a rugulosin-producing needle endophyte in two experiments. They were allowed to grow to sixth instar when survivors were collected. First, by using 3-yr old trees, a comparison was made of budworm growth on infected and uninfected trees. A second experiment used 4-yr old trees to study the effect of rugulosin content in the needle on growth. This permitted an examination of a dose response in relation to growth, and allowed us to eliminate the potential for differences in environment or foliar chemistry affecting the results. At sixth instar, budworms feeding on infected trees that contained rugulosin were smaller than those on uninfected trees. At needle concentrations above the dietary low observed effect level of rugulosin for C. fumerana >0.5 μg g⁻¹, a dose response was seen. For the first time, this demonstrates an inverse effect in outdoor nursery experiments between budworm weight and rugulosin concentration.     

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.     

Kairomone-mediated host finding by spruce budworm egg parasite,Trichogramma minutum

The host-finding responses of the egg parasite,Trichogramma minutum Riley, reared from its natural host,Choristoneura fumiferana (Clemens) and a laboratory host,Sitotroga cerealella (Oliver), to hexane extracts of scales of these hosts were compared in laboratory experiments. When presented withC. fumiferana eggs on filter papers treated with hexane and hexane extract of grain moth scales, egg finding byT. minutum was random. When presented with budworm eggs on filter papers treated with hexane and hexane extract of budworm scales,T. minutum found significantly more eggs on extract-treated filter papers, indicating the presence of kairomones on the scales of spruce budworms.T. minutum responded to kairomone-treated patches with a localized search behavior characterized by intense antennal tapping, decreased locomotion and increased turning within the patch, and repeated returns to the patch. In the absence of successful host finding, this response slowly waned. No significant difference in behavior ofT. minutum reared on eitherC. fumiferana orS. cerealella was observed.     

Some solid-phase syntheses of the sex attractant of the spruce budworm—trans-11-tetradecenal

The sex attractant of the spruce budworm,Choristoneura fumiferana,trans-11-tetradecenal (I), was synthesized on solid phases by three different routes from inexpensive 1,10-decanediol. The preparation of thetrans-alkene I directly on the polymer support was not completely stereoselective.The Use of Polymer Supports in Organic Synthesis, XIII.     

Sex Pheromone of the Common Sheep Moth, Hemileuca eglanterina, from the San Gabriel Mountains of California

The sex pheromone of Hemileuca eglanterina from the San Gabriel Mountains, California, was determined to be a combination of E10,Z12-hexadeca-10,12-dien-1-yl acetate, E10,Z12-hexadeca-10,12-dien-1-ol, and E10,Z12-hexadeca-10,12-dienal. Ratios of the compounds in extracts of female pheromone glands varied around a mean of 100 : 48 : 1.1 of the acetate, alcohol, and aldehyde, respectively. Field trials with synthetic compounds indicated that the optimum ratio of alcohol to aldehyde was 10 : 1 and that this ratio was more critical than the ratio of either compound to the acetate. A synthetic blend of 100 : 10 : 1 acetatendash;alcoholndash;aldehyde was effective at attracting male moths in the field. Additional compounds found in both extract and aeration samples failed to significantly increase trap catches of male moths, although some of these minor components elicited responses from male moth antennae in coupled gas chromatography–electroantennography studies.     

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.     

Effect of sex-pheromone concentration on behavior of three strains of western spruce budworm male moths

The responses of male western spruce budworm moths,Choristoneura occidentalis Freeman, to a range of concentrations of the major sex pheromone, 928 (E/Z)-11-tetradecenal (Ald), in polyvinyl chloride lures, were observed using the electroantennogram technique, a flight tunnel, and field-trapping bioassays. The responses to virgin female moths were also observed in the flight tunnel and field bioassays. The moths were from three strains: a nondiapausing laboratory colony; field-collected wild budworm; and laboratory-wild crosses. The mean peak amplitude of antennal response and the time required for the electroantennogram signal to return to the baseline after stimulation (lag) increased with Ald concentration in both laboratory and wild moths. However, at Ald concentrations of 0.005% and greater, the lag period of the wild male's antennae was significantly shorter than that of the laboratory male's. The mean number of moths caught in the field in delta sticky traps increased with Ald concentration, but the number of moths caught per trap was not significantly different between concentrations of 0.005 % and 0.5 %. The threshold concentration required to elicit upwind flight in the flight tunnel was between 0.0005 and 0.005% Ald; peak response occurred to 0.05 % Ald but was not significantly different from that to 0.005 % or 0.5% Ald. Moths from all three populations significantly reduced their net upwind groundspeed as they approached the pheromone lure. When pheromone concentration was increased, the net upwind groundspeed of laboratory and lab-wild moths, but not wild moths, was significantly reduced between 2 m and 1 m downwind from the pheromone lure. The three populations of moths differed significantly in the percentage of wing-fanning and copulatory attempts, and in the net upwind groundspeed of flight from 2 m to 1 m downwind from the lure.     

Roles of minor components in pheromone-mediated behavior of western spruce budworm male moths

The behavior of male western spruce budworm moths,Choristoneura occidentalis Freeman, was observed in a flight tunnel in response to virgin females and synthetic sex pheromone components, alone and in blends. Pheromone blends were also compared in the field using sticky trap bioassays. Pheromones were incorporated into small rods of polyvinyl chloride. The blend of 928 (E/Z)-11-tetradecenal-8911 (E/Z)-11-tetradecenyl acetate-8515 (E/Z)-11-tetradecenol (AldAcOH) that approximated that released from a virgin female moth elicited levels of response similar to those elicited by the female. This blend induced a significantly greater percentage of moths to fly upwind and land at the lure than did the Ald lure. In contrast to the flight-tunnel bioassays, the numbers of moths caught in Ald-baited sticky traps in the field were not significantly increased by the addition of Ac and OH lures. The net upwind groundspeed of flight in response to the 0.05% Ald lure was lower than that in response to the virgin females and was significantly increased by the addition of Ac + OH lures in two of three bioassays. The flight-tunnel bioassays support the hypothesis that the natural blend of major (Ald) and minor (Ac + OH) components stimulates the precopulatory behavior of western spruce budworm male moths at long range (> 1 m downwind) as well as at close range.     

Development of a bioluminescence assay for aldehyde pheromones of insects

A newly developed bioluminescent assay was used to measure quantitatively the amount of (E)-11-tetradecenal, the major component of the sex pheromone of the spruce budworm, trapped on Porapak Q^®. The bioluminescent response was linearly related to the amount of aldehyde either deposited on the absorbent or trapped from an airstream. However, the recovery of pheromone from Porapak was dependent on whether the air was prefiltered (through Porapak) or taken directly from the atmosphere. Furthermore, pheromone on Porapak was lost with time during the flow of air through the absorbent, indicating that trapping of aldehyde pheromone should be conducted for short periods of time for optimal recoveries. The applicability of the assay system for the rapid and direct measurement of the release rates of aldehyde pheromone lures was demonstrated for pheromone lures used for baiting spruce budworm traps.This investigation was supported by the U.S.D.A. Forest Service Program entitled Canada United States Spruce Budworms Program (23–345) and the Medical Research Council of Canada (MT-4314).     

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.     

Reinvestigation confirms action of Δ11-desaturases in spruce budworm moth sex pheromone biosynthesis

The biosynthesis of a large number of sex pheromone components of various moth species has been shown to start with common fatty acids and involve chain shortening by two carbons and introduction of a double bond at the 11–12 position. A recent report indicates that one of these common components, (E)-11-tetradecenyl acetate, is present in the eastern spruce budworm,Choristoneurafumiferana, but is not made by this pathway. Reinvestigation of this insect using in vivo and in vitro techniques indicates that the acetate indeed is made by a sequence of reactions similar to that used in other leafroller moths. In fact, evidence was found for the presence of several 11-desaturase systems in spruce budworm. One produced a large quantity of (Z)-11-hexadecanoic acid, and another produced (E)-11-tetradecanoic acid. It is not known if the small amount of (Z)-11-tetradecanoic acid is produced by either of those two systems or by a third system. A comparison with other species showed that cabbage looper moths have only the first system, redbanded leafroller moths use the last two systems, and European corn borer moths have all three.     

Behavioral responses of the female eastern spruce budwormChoristoneura fumiferana (Lepidoptera,Tortricidae) to the sex pheromone of her own species

Female eastern spruce budworm moths respond to the synthetic sex pheromone of their own species (a mixture ofcis- andtrans-11-tetradecenal) by walking, antennal grooming, flexation of the body, extension of the ovipositors, and oviposition. The sex pheromone is perceived by receptors on the antennae. Electroantennogram responses from the female are approximately two-thirds the amplitude of those obtained from males.     

Attraction of various tortricine moths to blends containingcis-11-tetradecenal

Blends containingcis-11-tetradecenal with the trans isomer orcis-11-tetradecen-1-ol were field tested against the tortricine speciesChoristoneura conflictana andCroesia semipurpurana. The results indicate thatcis-11-tetradecenal alone is a very potent attractant for maleC. conflictana whereas maleCroesia semipurpurana were strongly attracted to blends oftrans/cis-11-tetradecenal in the range 8020–90 10.Cenopis acerivorana andChoristoneura fumiferana were also caught with mixtures of 11-tetradecenal isomers whileChoristoneura fractivittana was very strongly attracted to the mixture ofcis-11-tetradecen-1-ol/cis-11-tetradecenal in the ratio 95/5.     

Identification of male cabbage looper sex pheromone attractive to females

A multicomponent pheromone produced by male cabbage looper moths that is attractive to female moths in a flight tunnel bioassay was isolated and identified. Based on analyses of hairpencil extracts of male cabbage loopers and volatiles emitted by males, the pheromone has been identified as a blend consisting of (S)-(+)-linalool,p-cresol, andm-cresol. The chirality of the major component, (S)-(+)-linalool, is important for behavioral response of females. These pheromonal compounds were also identified as volatiles released by males when males were exposed to the principal pheromone component of female cabbage loopers, (Z)-7-dodecen-1-ol acetate. The amount of male pheromone released was increased significantly when males were exposed to a combination of (Z)-7-dodecen-1-ol acetate and the odor from cabbage. Neither linalool nor the cresols were detected in volatiles from cabbage or from males exposed to cabbage odor.     

Repellency of oral exudate to eastern and western spruce budworm larvae (Lepidoptera: Tortricidae)

A two-choice feeding bioassay was used to investigate the intra-specific repellency of the larval oral exudate of eastern and western spruce budworms,Choristoneura fumiferana (Clem.) andC. occidentalis Free., respectively. Results of the bioassay indicated that feeding behavior on artificial diet-drop feeding stations was modified in the presence of exudate, with feeding stations treated with conspecific exudate being avoided when an untreated station was available 3 cm away. Feeding was suppressed when a single, exudate-treated station was provided, or when the treated and untreated stations were separated by only 1 cm. The repellent effect functioned both inter- and intraspecifically. When induced to produce exudate,C. occidentalis larvae were not immediately repelled by either their own or other individuals' exudate. However, 24 hr after induction, test larvae were repelled by exudate from either source. In both species, larval oral exudate probably functions to repel conspecific competitors.     

(2S,12Z)-2-Acetoxy-12-heptadecene: Major Sex Pheromone Component of Pistachio Twig Borer, Kermania pistaciella

The sex pheromone of the pistachio twig borer, Kermania pistaciella (Lepidoptera: Oinophilidae), one of the most important insect pests of pistachio, Pistacia vera, in Turkey and Iran, was identified. In gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectrometric analyses of pheromone gland extracts of female K. pistaciella from Turkey, (2S,12Z)-2-acetoxy-12-heptadecene was identified as the major candidate pheromone component. In field experiments in Turkey, lures containing synthetic (2S,12Z)-2-acetoxy-12-heptadecene attracted large numbers of male moths. Its attractiveness was significantly reduced by the presence of the R-enantiomer or of either enantiomer of the corresponding alcohol. (2S,12Z)-2-Acetoxy-12-heptadecene is the first pheromone component identified in the Oinophilidae and the first secondary acetate pheromone component identified in the Lepidoptera. An erratum to this article can be found at     

The role of monoterpenes in resistance of Douglas fir to western spruce budworm defoliation

We conducted defoliation experiments with 7- to 8-year-old clones of Douglas fir [Pseudotsuga menziesii (Mirb.) Franco var. glauca] to assess the role of monoterpenes as a resistance mechanism to western spruce budworm (Choristoneura occidentalis Freeman) defoliation. The grafted clones were derived from mature trees that showed resistance or susceptibility to budworm defoliation in the forest. All clones were exposed to either budworm defoliation or nondefoliation treatments in 1998 and 1999 under greenhouse conditions. We found that the total concentration of monoterpenes in current-year foliage varied greatly between two consecutive years in clones in the greenhouse and in their corresponding mature trees in the forest. Fractional composition of different monoterpenes was similar between different years and between clones and mature trees, indicating genetic control of this trait. Two different defoliation experiments were conducted to assess the importance of budburst phenology as a factor determining host plant resistance. In the 1998 experiment, budworm feeding was matched to the budburst of each individual plant. Monoterpene concentration was high in 1998, and budworm potential fitness was greater on clones from the resistant mature trees that had lower concentrations of total monoterpenes. In the 1999 experiment, budworm feeding was matched to budburst of the whole population of plants in order to mimic conditions similar to insects feeding on trees in the field. The concentration of monoterpenes was low in 1999, and budworm fitness was not related to monoterpenes. Total monoterpene concentration was negatively related to foliar nitrogen concentration, suggesting that C/N balance may affect monoterpene synthesis in needles. However, tree growth was not related to total monoterpene concentration. We concluded that expression of differences in budworm resistance among Douglas fir genotypes might be caused by interactions among multiple resistance mechanisms such as needle monoterpenes and tree budburst phenology.     

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     

Feeding responses of eastern spruce budworm larvae to sucrose and other carbohydrates

Spruce budworm larvae reared on an artificial diet were tested for preference of twelve carbohydrates over a water control. A strong preference for sucrose was seen, followed by fructose, inositol, and glucose. Male and female larvae do not differ in their responses to sucrose. The behavioral threshold for sucrose is at 10^–4 to 10^–3 M, with a peak response in the range of 0.01 to 0.05 M sucrose. Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae).Work leading to this publication was funded in part by a USDA Forest Service sponsored program entitled Canada/United States Spruce Budworms Program (grant 23–352), and in part by the National Sciences and Engineering Council of Canada (operating grant A9723), and in part by the Whitehall Foundation.