Correlation of housefly sex pheromone production with ovarian development

Pheromone production in the housefly was monitored during oögenesis and in ovariectomized insects by gas-liquid chromatography (GLC) and radio-GLC. The presence of vitellogenic ovaries was required for the initiation of (Z)-9-tricosene (muscalure), (Z)-9,10-epoxytricosane and (Z)-14-tricosen-10-one synthesis. Methylalkane synthesis was enhanced by developing ovaries. Insects ovariectomized within 12 hr after emergence produced no detectable amounts of (Z)-9-tricosene, C₂₃ epoxide nor C₂₃ ketone and synthesized less methylalkanes than the controls. This effect was reversed by ovary implants. When flies were ovariectomized after oviposition, synthesis of (Z)-9-tricosene, C₂₃ epoxide and C₂₃ ketone continued. Thus, initiation of the synthesis of these C₂₃ pheromone compounds required a vitellogenic ovary, but the ovary was not required to maintain synthesis.     

Female sex pheromone of Sesamia cretica: chemical and behavioural evidence for a three-component blend

The female‐produced sex pheromone of the Durra stem borer, Sesamia cretica (Lederer) (Lepidoptera: Noctuidae), had been previously characterized as a 75:25 blend of (Z)‐9‐tetradecenol (Z9‐14:OH) and (Z)‐9‐tetradecenyl acetate (Z9‐14:OAc) based on field trapping experiments. The low attraction of this blend in the field led us to further investigate the sex pheromone of this pest. Coupled gas chromatography with electroantennographic detection (GC‐EAD) analysis of female pheromone gland extracts consistently revealed three EAD‐active compounds. According to their GC retention times, mass spectra, and comparative EAG analyses with authentic standards, two of these compounds were found to be the previously reported components Z9‐14:OH and Z9‐14:OAc, whereas a third compound was identified as (Z)‐11‐hexadecenol (Z11‐16:OH). In wind tunnel experiments, the highest male responses were elicited by ratios of Z9‐14:OH, Z9‐14:OAc, and Z11‐16:OH, ranging from 90:1:9 to 90:5:5. In field tests, the 90:1:9 ratio of the blend loaded onto rubber septum dispensers was significantly more effective than single‐component, two‐component, and any other ratio of the three‐component blend. The greater effectiveness of this blend resulted in a more accurate detection of S. cretica flight activity compared with the previously reported two‐component blend.     

Electrophysiological and behavioural evidence for a fourth sex pheromone component in the turnip moth, Agrotis segetum

Abstract. In addition to the pheromone components (Z)-5-decenyl, (Z)-7-dodecenyl and (Z)-9-tetradecenyl acetate (Z5-10:OAc, Z7-12:OAc and Z9-14:OAc), it has previously been shown that the sex pheromone gland of the turnip moth, Agrotis segetum (Lepidoptera: Noctuidae Schiff) contains 10:OAc, 12:OAc, Z5-12:OAc, Z9-12:OAc, 11–12:OAc, Z5-14:OAc, Z7-14:OAc and Z11-16:OAc. To find out whether any of these additional compounds is involved in the sex pheromone communication in A. segetum, a comprehensive electro-physiological and behavioural investigation was conducted. Single-sensillum recordings on male antennae revealed three subtypes of sensilla among the previously so-called Z5-10:OAc sensilla. One subtype was identified having one receptor neurone (A) that responded to Z5-10:OAc with a large spike amplitude and another neurone (B) that responded to (Z)-5-decenol (Z5-10:OH) with a small spike amplitude. In another subtype the B neurone responded to Z5-12:OAc and sometimes also to 27-12:OAc and 10:OAc, in addition to responding to Z5-10:OH. In a third subtype the A neurone responded to all acetates identified from the female pheromone gland, whereas the small spike amplitude neurone was tuned to Z5-10:OH. A flight tunnel assay showed that blends composed of nine, eight or seven compounds were equivalent to the previously identified three-component pheromone blend in eliciting male behavioural responses. In field trapping tests, blends of eleven, nine or seven compounds did, however, catch significantly more moths than the three-component blend. Further assays showed that only 25- 12:OAc could significantly increase the catch numbers when added to the three-component blend, and thus qualified as a fourth pheromone component in A. segerum. The behavioural significance of additional female-produced acetates — for which males possess antennal receptors — is suggested, but may be impossible to confirm because of ‘diminishing returns’ when trying to refine a multicomponent pheromone further.     

Attractiveness of year-old polyethylene Isonet sex pheromone dispensers for Lobesia botrana

In this study, we have compared the release of sex pheromone from mating disruption dispensers exposed in the field for 12 months and from calling females. The main pheromone component of the grapevine moth, Lobesia botrana (D. and S.) (Lepidoptera: Tortricidae), is (E)‐7,(Z)‐9‐dodecadienyl acetate, and a minor component is (Z)‐9‐dodecenyl acetate. Aged dispensers from two different years emitted a much higher amount of both pheromone components than calling females. However, the summer temperature during field exposure influenced the release from mating disruption dispensers the following year. In the wind tunnel, male L. botrana were equally attracted to 12‐month, field‐exposed dispensers, a standard monitoring pheromone lure, and to synthetic (E)‐7,(Z)‐9‐dodecadienyl acetate sprayed at the rate of 0.6–60 ng h^?1. Field trapping tests confirmed that aged dispensers from both years were at least as attractive to L. botrana males as a standard monitoring pheromone lure. The possible contribution of previously applied dispensers to the mating disruption efficacy during following applications is discussed.     

Field evaluation of female sex pheromone components of the cotton bollworm,Heliothis armigera

(Z)-11-hexadecenal and (Z)-9-hexadecenal were ineffective lures forH. armigera males unless combined. Attraction depended upon perception of a 90%–99% combination of (Z)-11-hexadecenal with 1%–10% (Z)-9-hexadecenal. Increasing the level of (Z)-9-hexadecenal in the mixture to 26.2% reduced catches. Adding 2.3% (Z)-7-hexadecenal to the mixture did not enhance or reduce attraction, while adding 8.7% (Z)-11-hexadecenol significantly reduced male catches. The combination of (Z)-11-hexadecenal and (Z)-9-hexadecenal was effective only when released from rubber dispensers but not from polyethylene vials. A load of 2 mg of the mixture on rubber dispensers effectively attracted males for at least 31 days. TheH. zea lure which contained all the pheromonal components of that species was also effective in attractingH. armigera males. TheH. virescens lure attracted significantly fewerH. armigera males than theH. zea lure.

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Résumé Les (Z)-11-hexadecenal et (Z)-9-hexadecenal sont des attractifs sexuels in efficaces pour les mâles deH. armigera. L'attraction dépend de la perception d'un mélange de 90 à 99% de (Z)-11-hexadecenal avec 1 à 10% (Z)-9-hexadecenal, L'augmentation jusqu'à 26,2% de la teneur en (Z)-9-hexadecenal réduit les captures. L'addition de 2,3% de (Z)-7-hexadecenal au mélange ne modifie pas l'attractivité, tandis que celle de 8,7% de (Z)-11-hexadecenol, réduit significativement les captures de mâles. Le mêlange de (Z)-11-hexadecenal et de (Z)-9-hexadecenal n'a été efficace qu'avec des diffuseurs en caoutchouc, par contre il a été sans effet à partir de récipients de polyéthylène. Une charge de 2 mg de mélange dans des diffuseurs en caoutchouc attire effectivement les mâles pendant ou moins 31 jours. L'attractif sexuel deH. zea qui contient tous les constituants de la phéromone de cette espèce attire aussi efficacement les mâles deH. armigera. Celui deH. virescens attire significativement moins de mâles deH. armigera que l'attractif sexuel deH. zea.

     

Components of the sex pheromone of the currant pug moth, Eupithecia assimilata, a re-emergent hop pest in UK

After an absence of 50 years, the currant pug moth, Eupithecia assimilata Doubleday (Lepidoptera: Geometridae), has reappeared as an important pest of hops, Humulus lupulus L. (Cannabaceae), in the UK. Pheromone gland extracts from virgin female E. assimilata moths were shown to contain (3Z,6Z)‐cis‐9,10‐epoxyheneicosadiene (3Z,6Z‐cis‐9,10‐epoxy‐21:H) by gas chromatography linked to mass spectrometry. (3Z,6Z,9Z)‐heneicosatriene (3Z,6Z,9Z‐21:H) was also found as a minor component in one of two extracts. In field experiments, significant numbers of male E. assimilata moths were caught in traps baited with the (9S,10R)‐enantiomer of 3Z,6Z‐cis‐9,10‐epoxy‐21:H but not in those baited with the (9R,10S)‐enantiomer or racemic mixture. Addition of 3Z,6Z,9Z‐21:H at the ratio present in gland extracts greatly reduced the attractiveness of the epoxydiene.     

Sex ratio selection and multi-factorial sex determination in the housefly: a dynamic model

Sex determining (SD) mechanisms are highly variable between different taxonomic groups and appear to change relatively quickly during evolution. Sex ratio selection could be a dominant force causing such changes. We investigate theoretically the effect of sex ratio selection on the dynamics of a multi-factorial SD system. The system considered resembles the naturally occurring three-locus system of the housefly, which allows for male heterogamety, female heterogamety and a variety of other mechanisms. Sex ratio selection is modelled by assuming cost differences in the production of sons and daughters, a scenario leading to a strong sex ratio bias in the absence of constraints imposed by the mechanism of sex determination. We show that, despite of the presumed flexibility of the SD system considered, equilibrium sex ratios never deviate strongly from 1 : 1. Even if daughters are very costly, a male-biased sex ratio can never evolve. If sons are more costly, sex ratio can be slightly female biased but even in case of large cost differences the bias is very small (<10% from 1 : 1). Sex ratio selection can lead to a shift in the SD mechanism, but cannot be the sole cause of complete switches from one SD system to another. In fact, more than one locus remains polymorphic at equilibrium. We discuss our results in the context of evolution of the variable SD mechanism found in natural housefly populations.     

An evaluation of (Z)-9-tricosene and food odours for attracting house flies, Musca domestica, to baited targets in deep-pit poultry units

Field trials investigating the effect of food baits on catches of Musca domestica at toxic targets impregnated with the female sex pheromone, (Z)-9-tricosene, were conducted in a caged-layer deep-pit poultry unit in southern England. Targets treated with an Alfacron-sugar mixture and baited with 2.5 g of 40% (Z)-9-tricosene beads caught significantly greater numbers of both male and female M. domestica than control targets. Egg and milk-baited targets were less attractive than controls, while brewers yeast slightly increased the numbers of M. domestica attracted. However, the inclusion of brewers yeast in (Z)-9-tricosene-impregnated targets produced a significant reduction in the number of male M. domestica attracted. Increased female attraction was elicited by baiting the targets with 2-phenylethanol, at the quantities of 1 mg and 10 mg. However, 2-phenylethanol had no effect on female attraction when presented in conjunction with (Z)-9-tricosene. The implications of these results in relation to the control of M. domestica populations in poultry units are discussed.     

淡剑袭夜蛾性信息素活性成分的鉴定和田间诱蛾研究

提取和鉴定了淡剑袭夜蛾(Sidemia depravata)的性信息素主要组分,并进行了田问试验．结果表明,雌蛾性腺体提取物中主要性信息素组分为顺-9-十四碳烯醇醋酸酯(29-14：Ac)和顺-9,反-12-十四碳烯醇醋酸酯(Z9E12-14：Ac)。在这2种化合物配制的二元组份诱芯中,以Z9-14：Ac／Z9E12-14：Ac＝5／5和6／4比例配制成的诱芯诱蛾效果较好,田间诱蛾量显著高于以1：9、2：8和3：7比例配制的诱芯。     

Female sex pheromone components of Helicoverpa gelotopoeon: first heliothine pheromone without (Z)-11-hexadecenal

Helicoverpa gelotopoeon Dyar is a very important pest of economic importance on cotton in Argentina. Analysis of female pheromone gland extracts prepared from 1‐ to 2‐day‐old virgin female moths demonstrated the presence of a 1 : 0.84 blend of hexadecanal (16:Ald) and (Z)‐9‐hexadecenal (Z9‐16:Ald), with trace quantities of tetradecanal in some samples, 2.4% of 16:Ald. The average quantity of Z9‐16:Ald extracted per female was estimated to be 33 ng, with a range of 18.9–46.4 ng per female when collected 2–3 h into the scotophase. In field trials conducted in both cotton and tomato crops in Santiago del Estero, Argentina 1 : 1 blends of 16:Ald and Z9‐16:Ald caught significantly more male H. gelotopoeon than Z9‐16:Ald alone, although there was no significant difference between blends containing between a 0.2 : 1 and 2 : 1 ratio of 16:Ald and Z9‐16:Ald. There was no analytical evidence for the presence of (Z)‐11‐hexadecenal (Z11‐16:Ald) in pheromone gland extracts, although this compound has been identified in all female sex pheromones of Heliothinae to date. In field trials, the addition of Z11‐16:Ald at the 1% level to either a 1 : 1 blend of 16:Ald and Z9‐16:Ald or Z9‐16:Ald alone significantly reduced the catch of male H. gelotopoeon. Sympatric Heliothis virescens were not caught in any of the blends tested for H. gelotopoeon, but were caught in low numbers in traps baited with a 4 : 100 blend of (Z)‐9‐tetradecenal and (Z)‐11‐hexadecenal.     

Canalization of the redbanded leafroller moth sex pheromone blend

Previous attempts to shift the (Z)-11-/(E)-11-tetradecenyl acetate ratio of pheromone components in the redbanded leafroller moth (RBLR), Argyrotaenia velutinana, by several selection protocols showed that this ratio is strongly canalized. Analysis of the complete seven-component blend, however, showed that a Geneva laboratory stock of RBLR had a lower percent (20%) of the E9–12:OAc minor component compared to the E11–14:OAc component than a population of RBLR from North Carolina (31%). Hybrid populations from these two cultures were used in a two-way family truncation selection scheme in which families were selected for either the lowest (low line) or the highest (high line) ratio of E9–12:OAc/E11–14:OAc. After three generations of selection, the low line had 14% E9–12:OAc relative to E11–14:OAc and the high line had 42%. The selection pressure was removed in generations 4–9, and the low line remained unchanged at 14% E9–12:OAc; but in the high line, it drifted to 53%. Studies were conducted to estimate heritability and realized heritability. The realized heritability calculated for each generation of selection averaged 1.14 for the low line and 1.50 for the high line. These calculations, along with estimated heritability values of 0.416 and 0.644 for reciprocal crosses, indicate some plasticity in the E9–12:OAc/E11–14:OAc ratio. This ratio was positively correlated to the total amount of 12-carbon components to 14-carbon components, but was negatively correlated to the Z/E ratio of Δ11-tetradecenylacetates. The results of two studies on the canalization of various components of the RBLR sex pheromone blend indicate that there is limited potential in this insect for manipulation of the blend ratios in the laboratory.     

Field evaluation of female sex pheromone of the artichoke moth, Gortyna xanthenes

Field responses of Gortyna xanthenes (Germar) males to traps baited with different mixtures of the female sex pheromone components were evaluated in an artichoke field. Catches were compared to those obtained by virgin females and light traps. The best results were achieved by utilizing a mixture of 5 mg Z11-16:Ald + 0.15 mg Z9-16:Ald + 0.15 mg 16:Ald + 0.12 mg Z11-16:OH, which captured only G. xanthenes males 3–5 times more than light traps and ca two times more than one virgin female. Starting from the basic mixture Z11-16:Ald (89–92%) + Z9-16:Ald (2–4%), the addition of 16:Ald (2–4%) and of Z11-16:OH (2–3%) produced an increase of G. xanthenes and a decrease of H, armigera (Hb.) catches. The inhibitory action of Z11-16:OH towards H. armigera males was confirmed.

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Evaluation sur le terrain des constituants de la phéromone sexuelle de la noctuelle de l'artichaut (Gortyna xanthenes)

Résumé Dans un champ d'artichaut de la province de Salerno on a evalué la réponse des mâles de G. xanthenes à mélanges divers des constituants de la phéromone sexuelle produite par la femelle. Les meilleurs résultats ont été obtenus en employant un mélange de 5 mg Z11-16:Ald + 0.15 mg Z9-16:Ald + 0.15 mg 16:Ald + 0.12 mg Z11-16:OH, qui a capturé seulement les mâles de G. xanthenes en mesure 3–5 fois plus élevée que les pièges lumineux et ca. deux fois plus élevée que la femelle vierge. En partant du mélange base Z11-16:Ald (89–92%) + Z9-16:Ald (2–4%), l'addition de 16:Ald (2–4%) et de Z11-16:OH (2–3%) a causé un accroissement des captures de G. xanthenes et une diminution des captures de Heliothis armigera. Cela confirme l'action inhibitoire du Z11-16:OH à l'égard des males de H. armigera.

     

A two-component sex attractant for males of the geometrid moth Alsophila quadripunctata

The binary mixture of the hydrocarbons (3Z,6Z,9Z)-3,6,9-nonadecatriene and (6Z,9Z)-6,9-nonadecadiene is a sex attractant for males of Alsophila quadripunctata Esp. (Lepidoptera, Geometridae). Triene/diene ratios from 1:1 to 1:0.01 attracted males in the field, the ratios 1:1 and 1:0.7 were most effective. The triene on its own also attracted males, but in significantly lower numbers than the 1:1 mixture. In a dosage test the highest dose, 1800 g of 1:0.8 [triene:diene] mixture applied to rubber caps was most effective. This is the first report of a geometrid sex attractant with field activity containing more than one component.

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Zusammenfassung Die binäre Mischung der Kohlenwasserstoffe (3Z,6Z,9Z)-3,6,9-Nonadecatrien [ZZZ3,6,9-19:H] und (6Z,9Z)-6,9-Nonadecadien [ZZ6,9-19:H] in den Verhältnissen 1:1 bis 1:0.01 erwies sich im Freiland als Sexuallockstoff für Männchen von Alsophila quadripunctata (Geometridae, Lepidoptera). Dabei besaß die 1:1-Mischung die beste Fängigkeit, das Nonadecatrien allein lockte schlechter, während das C₁₉-Alkadien unwirksam war. Ein Dosiswirksamkeitstest ergab bei Köderbeladungen von 1.8 mg (Trien:Dien + 1:0.8) die besten Fangergebnisse. Im Verlauf der Untersuchungen wurden weitere zwölf geometrische-oder positionsisomere C₁₉-Polyenkohlenwasserstoffe getestet, wobei keine der geprüften Verbindungen Attraktant-oder Inhibitorwirkung zeigte.

     

Optimization of pheromone lure and trap design for monitoring the fir coneworm, Dioryctria abietivorella

The major components of the sex pheromone of Dioryctria abietivorella (Groté) (Lepidoptera: Pyralidae) were recently identified as (9Z,11E)‐tetradecadien‐1‐yl acetate (9Z,11E‐14:Ac) and a polyunsaturated, long‐chain hydrocarbon (3Z,6Z,9Z,12Z,15Z)‐pentacosapentaene (C25 pentaene). The optimal ratio of these components and the role of potential minor components were not fully determined in the initial short report on the pheromone's identification. We tested different ratios of the two major components loaded into grey halobutyl rubber septum dispensers, placed in sticky traps deployed in conifer breeding arboreta. The optimal ratio of the two components was 200 µg 9Z,11E‐14:Ac to 2000 µg C25 pentaene. (Z)‐9‐Tetradecen‐1‐yl acetate, which had been identified previously in female pheromone gland extracts, and five other potential minor pheromone components, were tested individually as additions to the optimized two‐component lure blend. None of the ternary blends were more attractive than the optimized two‐component blend, at the ratios tested. Two lure adjuvants, a UV stabilizer (Sumisorb 300) and the antioxidant butylated hydroxytoluene, added individually or together, did not affect the attractiveness of the optimized lure blend. The Pherotech diamond sticky trap baited with the optimized lure blend was the most effective trap design among eight types of sticky trap and a bucket style trap that were tested. Traps baited with synthetic lures were as attractive as traps baited with virgin female moths. The optimized two‐component lure blend in the Pherotech diamond trap is recommended for monitoring fir coneworm infestations. The availability of an effective synthetic pheromone opens the possibility for control tactics using mating disruption or attract‐and‐kill techniques.     

Sex pheromone of the butterbur borer, Ostrinia zaguliaevi

The sex pheromone blend of the butterbur borer, Ostrinia zaguliaevi (Lepidoptera: Pyralidae) was analyzed by means of gas chromatography-electroantennographic detection (GC-EAD), GC-mass spectrometry and a series of wind-tunnel bioassays. Four EAD-active compounds were detected in the female sex pheromone gland extract, and these were identified as tetradecyl acetate (14:OAc), (Z)-9-tetradecenyl acetate (Z9-14:OAc), (E)-11-tetradecenyl acetate (E11-14:OAc) and (Z)-11-tetradecenyl acetate (Z11-14:OAc). The average amounts ± s.d. of the four compounds in a single sex pheromone gland were 7.9±3.7 ng, 10.1±3.2 ng, 1.1±0.5 ng and 11.6±5.1 ng, respectively. In a wind-tunnel bioassay, the ternary blend of Z9-, E11- and Z11-14:OAc at a ratio found in the sex pheromone gland (45:5:50) elicited the same behavioral responses from the males as did virgin females and pheromone gland extract. Removal of any single compound from the ternary blend significantly diminished the pheromonal activity, whereas addition of 14:OAc to the ternary blend had no effect on the males' behavioral responses. Therefore, it was concluded that the sex pheromone blend of O. zaguliaevi is composed of Z9-14:OAc, E11-14:OAc and Z11-14:OAc at a ratio of 45:5:50.     

Identification of the sex pheromone of Ostrinia palustralis

By means of gas chromatography with electroantennographic detection, gas chromatography-mass spectrometry and a series of bioassays, (E)-11-tetradecenyl acetate (E11-14:OAc) and (Z)-11-tetradecenyl acetate (Z11-14:OAc) at a ratio of 99:1 were identified as female sex pheromone components of Ostrinia palustralis. The average amounts of E11- 14:OAc and Z11-14:OAc in a single sex pheromone gland were 37.2±24.4 ng and 0.3±0.2 ng, respectively. In a wind-tunnel bioassay, the binary blend of E11- and Z11-14:OAc elicited the same male behavioral responses as did virgin females.     

The effects of laboratory culturing on (Z)-9-tricosene (muscalure) quantities on female houseflies

Using gas chromatography the relative amounts of (Z)-9-tricosene (muscalure) and some other hydrocarbons on the cuticle of 1- to 20-day-old houseflies (Musca domestica L.) from different strains were determined. Flies from a WHO strain, in culture since 1961, and first-generation laboratory-cultured flies from two wild-type strains from a poultry breeding and a cow-house with pigsty, respectively, were compared. On WHO females hydrocarbons with 23–25 C atoms constituted about 65% of the total hydrocarbons, whereas on wild-type females less than 2% of these compounds was present. (Z)-9-tricosene comprised up to 20–30% of the total hydrocarbons on 5- to 20-day-old WHO females, whereas less than 0.5% (Z)-9-tricosene was present on the wild-type females. We also compared the amounts of (Z)-9-tricosene and some other hydrocarbons on female houseflies, kept in culture in the laboratory for several generations. It appeared that whereas on first-generation wild-type females hardly or no (Z)-9-tricosene could be detected, the amounts of this substance had increased considerably after some tens of generations in the laboratory. It is suggested that this was due to selection in subsequent generations of high-density populations. Production of (Z)-9-tricosene and of tricosane was shown to be closely linked. Selection did not affect the production of other cuticular hydrocarbons by the females. It is suggested that in mixed populations (both sexes together in a cage) in the course of time (Z)-9-tricosene is transferred from females to males and (Z)-9-heptacosene from males to females. It is concluded that reproductive ability of houseflies does not primarily depend on the amounts of (Z)-9-tricosene on females, although higher amounts of this substance may increase contacts between males and females.     

Spectroscopic evidence for the formation of a transient species during cytochrome P-450scc induced hydroperoxysterol-glycol conversions

Spectroscopic analysis of the interaction of the epimeric 20-hydroperoxy derivatives of cholesterol with bovine adrenocortical cytochrome P-450scc preparations suggested the formation of a transient species. The intermediate was detected at 4 degrees C and characterized by a minimum at 412 nm in the difference spectrum.