Variation in sex pheromone composition in large fruit-tree tortrix Archips podana Scop. (Lepidoptera: Tortricidae)

Variation in sex pheromone composition was studied in large fruit-tree tortrix. The pheromone contained cis-11-tetradecenyl acetate and trans-11-tetradecenyl acetate as macrocomponents as well as microcomponents cis-11-tetradecenol and trans-11-tetradecenol with the prevalence of cis-11-tetradecenyl acetate and cis-11-tetradecenol. The pheromone composition depended in part on the genotype of the male parent. Inbreeding increased the proportion of trans-11-tetradecenyl acetate and trans-11-tetradecenol.     

Electroantennogram responses of the male moth, Argyrotaenia velutinana to mixtures of sex pheromone components of the female

Electroantennogram (EAG) responses were recorded from male redbanded leafroller moth, Argyrotaenia velutinana, antennae using mixtures of the three female-produced sex pheromone components: cis-11-tetradecenyl acetate, trans-11-tetradecenyl acetate, and dodecyl acetate (12:Ac). Binary mixtures containing 3%, 8%, and 15% trans in cis elicited significantly higher amplitude responses than other isomeric mixtures as well as pure cis and trans alone. The higher responses to such mixtures were less than additive at high dosages and additive at lower dosages. Receptor adaptation studies using the two isomers support a previous single unit study demonstrating the presence of at least two functionally different receptor sites on male A. velutinana antennae; adaptation to an airstream containing one isomer did not eliminate response to the opposite isomer presented concurrently. An airstream containing the third component, 12:Ac, caused a significant slowing of the recovery rate during the entire recovery period of EAG responses to cis but not trans, suggesting that a possible temporal modulation of neuronal response by 12:Ac may be a means of coding for this component by antennal sensory neurons.     

Electroantennogram responses of two tortricid moths using two-component sex pheromones

Antennal responses of male Adoxophyes orana and Clepsis spectrana to their two pheromone components, cis-9- and cis-11-tetradecenyl acetate (tda) were studied. Responses to a dilution series of the components alone and in five mixture ratios were recorded. The main pheromone component for A. orana (cis-9-tda) and for C. spectrana (cis-11-tda) gave the highest electroantennogram (EAG) responses, respectively, but mixtures corresponding to the optimum field attracting ratios for each species elicited higher responses than the separate components.The geometrical isomers, trans-9- and trans-11-tda, which inhibit male A. orana response in the field, do not appear to give an additive EAG response when mixed with the pheromone components.     

Sex pheromone components of the leafroller moth Pandemis cerasana

Female-tip washings of the leafroller moth, Pandemis cerasana, were found to contain (E)-11-tetradecenyl acetate, (Z)-11-tetradecenyl acetate, (E)-11-tetradecenol, (Z)-11-tetradecenol and tetradecyl acetate, based on chemical analysis and electroantennogram tests. The relative amounts of these compounds in the gland were ca. 64:21:10:3:2 in the order named. Only (E)-11 and (Z)-11-tetradecenyl acetate were required for attraction of males to trap dispensed in the ratio 3:1, respectively.     

Reidentification of pheromone composition of Sparganothis sulfureana (Clemens) and evidence of geographic variation in male responses from two US states

GC-EAD analyses of pheromone gland extracts of calling female Sparganothis sulfureana revealed at least 6 compounds that consistently elicited antennal responses from male antennae. In addition to the major pheromone compound, (E)-11-tetradecenyl acetate (E11–14:OAc), which was previously reported, the other compounds were found to be (E)-9-dodecenyl acetate (E9–12:OAc), (Z)-9-dodecenyl acetate (Z9–12:OAc), (Z)-9-tetradecenyl acetate (Z9–14:OAc), (Z)-11-tetradecenyl acetate (Z11–14:OAc), and (E)-11-tetradecenol (E11–14:OH). Tetradecyl acetate, hexadecyl acetate and hexadecenyl acetates were also present in the extracts, but elicited no EAG response from male antennae. Wind tunnel tests demonstrated that males from New Jersey responded equally well to a blend containing five pheromone components in relative to the pheromone glands of calling females. Different male-response profiles from field-trapping tests conducted in the states of Wisconsin and New Jersey were observed, respectively. Significantly higher numbers of male S. sulfureana were caught in New Jersey in traps baited with the binary blend of E11–14:OAc (30 μg) with 1% of Z11–14:OAc, but males from Wisconsin responded equally well to traps containing blends of E11–14:OAc with 0–10% of Z11–14:OAc. The addition of more than 10% of Z11–14:OAc to the primary pheromone compound reduced male captures significantly in both states. Male catches were doubled by adding E9–12:OAc and E11–14:OH to the most attractive binary blend in both states. The trapping test with caged live virgin female moths showed that males in Wisconsin preferred females from the local population than those from New Jersey. The differences in male responses observed may indicate the existence of pheromone polymorphism in this species.     

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.     

Electroantennogram response gradient on the antenna of the European corn borer, Ostrinia nubilalis

The electroantennogram (EAG) response of male moths of the European corn borer (Ostrinia nubilalis) to their two pheromone components, (Z)- and (E)-11-tetradecenyl acetates, were studied. The EAG amplitude of the fast initial phase was approximately proportional to the antennal length. Local EAG responses were obtained along the length of the antenna with multiple electrodes. Response-interaction studies indicated a unique spread of the EAG response in a proximal direction in the antenna. A possible mechanism of the EAG summation was considered. The EAG response characteristic was similar for both (Z)- and (E)-11-tetradecenyl acetate along the length of the antenna.     

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.     

Broadly and narrowly tuned odorant receptors are involved in female sex pheromone reception in Ostrinia moths

Mate-finding communication in many moths is mediated by sex pheromones produced by females. Since the differentiation of sex pheromones is often associated with speciation, it is intriguing to elucidate how the changes in sex pheromones are tracked by the pheromone recognition system of the males. Moths of the genus Ostrinia, which show distinct differentiation in female sex pheromones, are good models to study this. The present study was initiated with the aim of identifying ORs from Ostrinia scapulalis that respond to its own pheromone components, (E)-11- and (Z)-11-tetradecenyl acetates. We isolated six OR gene candidates (OscaOR3–8) from O. scapulalis. The same set of genes homologous to OscaOR3–8 were conserved in all (eight) Ostrinia species examined in addition to the previously reported OscaOR1 (tuned to (E)-11-tetradecenol) and the Or83b homologue OscaOR2. OscaOR3 not only responded to (E)-11- and (Z)-11-tetradecenyl acetates, but also to the pheromone components of the congeners, (Z)-9-, (E)-12-, and (Z)-12-tetradecenyl acetates. OscaOR4 responded with a relatively high specificity to (E)-11-tetradecenyl acetate. While OscaOR5 responded only marginally to a few pheromone components, OscaOR6–8 did not respond to any of the compounds tested. A few conserved ORs, including a unique one with very broad responsiveness, appear to be involved in the sex pheromone reception in O. scapulalis. The findings of the present study are discussed with reference to knowledge on electrophysiological response profiles of olfactory receptor neurons in Ostrinia moths.     

Notes on New Zealand mammals 10. Effect of observer position on group size and level of aggression of mating brown hares

Abstract

The sex pheromones of two endemic New Zealand tortricid species were investigated. Females of the first species, Cnephasia jactatana, a pest of kiwifruit, were found to produce (Z)-11-tetradecenyl acetate (Z11-14:OAc) in their sex pheromone gland. When tested in the field, this compound on its own elicited significant catches of male C. jactatana. However, the addition of even very small amounts (0.3%) of the geometric isomer, (E)-11-tetradecenyl acetate (E11-14:OAc), to Z 11-14:OAc virtually suppressed trap catches. The sex pheromone of the second species, Merophyas leucaniana, was found to consist of a mixture of Z11-14:OAc, E11-14:OAc, tetradecyl acetate, and (Z)-11-tetradecenol. This species is morphologically very similar to the congeneric species, M. divulsana, the Australian lucerne leafroller. The sex pheromone of M. leucaniana is also very similar to that of the so-called “Z-type” of M. divulsana, which is now found in New Zealand. The pheromone components of C. jactatana and M. leucaniana are discussed in relation to those found in other tortricids.     

Biosynthesis of the acetate ester precursor of the spruce budworm sex pheromone by an acetyl CoA: Fatty alcohol acetyltransferase

The biosynthesis of 11-tetradecenyl acetate, the major storage precursor of the aldehyde pheromone of Choristoneura fumiferana, the eastern spruce budworm, has been found to be catalyzed by an acetyl-CoA: fatty alcohol acetyltransferase. In vitro, acetyltransferase activity was found almost exclusively in extracts from the pheromone producing gland, and could be demonstrated in vivo by topical application of [¹⁴C]tetradecanol to the glands. Moreover, the activity was under developmental regulation, being low before and immediately after emergence of the moths from the pupal stage, and rising to a maximum in concert with the increase in glandular pheromone levels. Maximum activity with saturated alcohols was observed for acceptors of 12 to 15 carbons in chain length, with higher activities being found for the cis or trans monounsaturated analogs. The specificity of this enzyme with respect to substrate, morphological location and developmental regulation, indicates that it plays a key role in regulation of pheromone biosynthesis.     

Evidence of olfactory antagonistic imposition as a facilitator of evolutionary shifts in pheromone blend usage in Ostrinia spp. (Lepidoptera: Crambidae)

Olfactory receptor neuron (ORN) response was measured to assess why some males ("rare males") of the Asian corn borer (ACB), Ostrinia furnacalis, have a broad behavioral response to fly upwind to both the ACB and the European corn borer (ECB), Ostrinia nubilalis, pheromone blends. We performed single-sensillum electrophysiological recordings on ACB males that had been behaviorally assessed for upwind flight response to the ACB blend [60:40 (Z)-12-tetradecenyl acetate (Z12-14:OAc) to (E)-12-tetradecenyl acetate (E12-14:OAc)], as well as to ECB (Z-strain) and ECB (E-strain) blends [3:97 and 99:1 (Z)-11-tetradecenyl acetate (Z11-14:OAc) to (E)-11-tetradecenyl acetate (E11-14:OAc)]. Sensilla from all types of males had large- and small-spike-sized ORNs responding strongly to Z12- or E12-14:OAc, but weakly to Z11- and E11-14:OAc. In the majority of males ("normal males") that flew upwind only to the ACB blend, Z11-14:OAc elicited responses in an intermediate spike-sized ORN associated with behavioral antagonism that is mainly tuned to (Z)-9-tetradecenyl acetate (Z9-14:OAc). In the rare-type ACB males that flew to both the ACB and ECB pheromone blends, Z11-14:OAc did not stimulate this ORN. Increased responsiveness to ancestral pheromone components by ORNs associated with behavioral antagonism could be instrumental in reproductive character displacement, or in reinforcement and reproductive isolation during speciation by helping to increase assortative mating between males and females in derived populations that use novel sex pheromone blends.     

Altered olfactory receptor neuron responsiveness in rare Ostrinia nubilalis males attracted to the O. furnacalis pheromone blend

Three percent of E-strain Ostrinia nubilalis males fly upwind in response to the Ostrinia furnacalis pheromone blend [a 40:60 ratio of (E)-12-tetradecenyl acetate to (Z)-12-tetradecenyl acetate (E12-14:OAc to Z12-14:OAc)], in addition to their own pheromone blend [a 99:1 ratio of (E)-11-tetradecenyl acetate to (Z)-11-tetradecenyl acetate) (E11-14:OAc to Z11-14:OAc)]. We assessed the olfactory receptor neuron (ORN) responses of these behaviorally "rare" males versus those of normal males. For the three ORNs housed within each sensillum, we tested responsiveness to Z12-14:OAc, E12-14:OAc, Z11-14:OAc, E11-14:OAc, and the behavioral antagonist (Z)-9-tetradecenyl acetate (Z9-14:OAc). Z11-14:OAc, E11-14:OAc, and Z9-14:OAc stimulated ORNs exhibiting distinct small, large, and medium spike sizes, respectively. For rare and normal males, both Z12-14:OAc and E12-14:OAc usually elicited responses from the largest-spiking ORN. In many ORNs of normal males, Z12-14:OAc or E12-14:OAc stimulated the smaller-spiking ORN that is responsive to Z11-14:OAc. In rare males, detectable ORN responses from the smaller-spiking ORN in response to Z12- and E12-14:OAc were virtually non-existent. These differences in ORN tuning in rare males will tend to create an ORN firing ratio between the large- and small-spiking ORNs in response to the O. furnacalis blend that is similar to that elicited by the O. nubilalis blend.     

Effects of egg-to-adult development time and adult age on olfactory neuron response to semiochemicals in European corn borers

We used the cut-sensillum technique to assess the effect of both adult age and egg-to-adult development time on olfactory neuron responses of Z strain moths of the European corn borer, Ostrinia nubilalis. Compounds tested included the pheromone components, (Z)-11-tetradecenyl acetate and (E)-11-tetradecenyl acetate, the behavioral antagonist, (Z)-9-tetradecenyl acetate, and components of the O. furnicalis (Asian corn borer) sex pheromone, (Z)-12-tetradecenyl acetate and (E)-12-tetradecenyl acetate. The proportion of moths having neurons responding to the two O. nubilalis sex pheromone components and antagonist increased with longer development time and age. The spike frequency of neurons responding to (E)-11-tetradecenyl acetate and the antagonist increased with longer development time. Fourteen of 45 moths with neurons sensitive to either of the O. nubilalis pheromone components responded to (Z)-12-tetradecenyl acetate or (E)-12-tetradecenyl acetate. The likelihood of (Z)-12-tetradecenyl acetate stimulating a neuron similar in spike shape and waveform to that responding to (E)-11-tetradecenyl acetate increased with development time.     

Relative attractiveness of incomplete and complete blends of synthetic pheromone to male obliquebanded leafroller (Lepidoptera: Tortricidae) moths in a flight tunnel and in apple orchards: implications for sex pheromone-mediated mating disruption of this species

The relative attractiveness of synthetic Choristoneura rosaceana (Harris) pheromone consisting of the major compound Z-11-tetradecenyl acetate (Z11-14:OAc), or the major compound plus one, two, or all three of the minor compounds E-11-tetradecenyl acetate (E11-14:OAc), Z-11-tetradecenol (Z11-14:OH), and Z-11-tetradecenal (Z11-14:Ald), in amounts similar to those in the natural pheromone, was compared in a flight tunnel and in apple orchards. In the flight tunnel, there was an increase in the proportion of moths that were activated when E11-14:OAc was combined with the main compound, but no detectible additional increases in response with the addition of Z11-14:OH and Z11-14:Ald. The time required for activation was twice as long when using the major compound than when using the major compound and one or more of the minor compounds. There was a trend of increase in the proportion of males initiating the take-off, lock-on, close-in, and touchdown phases of upwind flight when progressively more complete blends were used. The time required for the initiation of these behavioral phases was statistically similar for each treatment. The proportion of moths landing at the pheromone source was 10 times greater when using the complete blend than when using the main compound alone. Similar results were obtained in apple orchards, where the four-compound blend was 17-55 times more attractive than the main compound. The superior attractiveness of the four-compound pheromone compared with the major compound, or to the major compound plus the minor compound E11-14:OAc, suggests that it would be the most effective synthetic pheromone if competitive attraction were an important mechanism of disruption in this species.     

Sex pheromone of the moth,Antheraea polyphemus

The sex pheromone system of Antheraea polyphemus was characterized from female abdominal tips by classical and electroantennogram techniques as rans-6,cis-11-hexadecadienyl acetate and trans-6,cis-11-hexadecadienal. A 90 : 10 mixture of acetate and aldehyde was highly attractive to wild males in the field. The synthetic pheromone and A. polyphemus females were not attractive to released Antheraea pernyi males.     

Identification of the Sex Pheromone of the Tree Infesting Cossid Moth Coryphodema tristis (Lepidoptera: Cossidae)

The cossid moth (Coryphodema tristis) has a broad range of native tree hosts in South Africa. The moth recently moved into non-native Eucalyptus plantations in South Africa, on which it now causes significant damage. Here we investigate the chemicals involved in pheromone communication between the sexes of this moth in order to better understand its ecology, and with a view to potentially develop management tools for it. In particular, we characterize female gland extracts and headspace samples through coupled gas chromatography electro-antennographic detection (GC-EAD) and two dimensional gas chromatography mass spectrometry (GCxGC-MS). Tentative identities of the potential pheromone compounds were confirmed by comparing both retention time and mass spectra with authentic standards. Two electrophysiologically active pheromone compounds, tetradecyl acetate (14:OAc) and Z9-tetradecenyl acetate (Z9-14:OAc) were identified from pheromone gland extracts, and an additional compound (Z9-14:OH) from headspace samples. We further determined dose response curves for the identified compounds and six other structurally similar compounds that are common to the order Cossidae. Male antennae showed superior sensitivity toward Z9-14:OAc, Z7-tetradecenyl acetate (Z7-14:OAc), E9-tetradecenyl acetate (E9-14:OAc), Z9-tetradecenol (Z9-14:OH) and Z9-tetradecenal (Z9-14:Ald) when compared to female antennae. While we could show electrophysiological responses to single pheromone compounds, behavioral attraction of males was dependent on the synergistic effect of at least two of these compounds. Signal specificity is shown to be gained through pheromone blends. A field trial showed that a significant number of males were caught only in traps baited with a combination of Z9-14:OAc (circa 95% of the ratio) and Z9-14:OH. Addition of 14:OAc to this mixture also improved the number of males caught, although not significantly. This study represents a major step towards developing a useful attractant to be used in management tools for C. tristis and contributes to the understanding of chemical communication and biology of this group of insects.     

Behavioural and electrophysiological activity of unsaturated analogues of the pheromone tetradecyl acetate in the small ermine moth Yponomeuta rorellus

ABSTRACT In field-trapping experiments the unsaturated analogues ( E )-6-, ( E )-12-, and ( Z )-12-tetradecenyl acetate were as attractive to male Yponomeuta rorellus Latr. as the native pheromone component tetradecyl acetate. All four analogues attracted more males than virgin females did, whereas ( Z )-6-, ( E )-11-, ( Z )-10- and ( Z )-11-tetradecenyl acetate were essentially non-attractive. Addition of 1–30% of ( Z )-11-tetradecenyl acetate to the pheromone tetradecyl acetate reduced the attraction to less than 2%. Flight tunnel experiments with Y. rorellus confirmed the activity of the ( E )-6- and ( E )-12-tetradecenyl acetates and demonstrated the activity of ( E )-7-tetradecenyl acetate as well. These analogues elicited orientation behaviour, upwind flight and landing at the odour source as frequently as the native pheromone did. Single sensillum recordings from male Y. rorellus showed two types of cells in most sensilla. A large spike amplitude cell was stimulated by tetradecyl acetate and the unsaturated analogues ( E )-11-, ( E )-6- and ( E )-12-tetradecenyl acetate, and to a lower extent by the ( Z )-6-, ( Z )-11- and ( Z )-12-isomers. A cell with medium spike amplitude was stimulated by ( Z )-9-tetradecenyl and ( Z )-11-hexadecenyl acetate. Some sensilla contained a third cell firing with a small spike amplitude which was activated by ( Z )-11-tetradecenol. Thus the tetradecyl acetate receptor was stimulated not only by the behaviourally active analogues, but also by behavioural antagonists. The interaction of ( E )-11-tetra-decenyl acetate and tetradecyl acetate with the same antennal receptor cell was also demonstrated in Y.cagnagellus . Electrophysiological discrimination between behavioural attractants and antagonists and the role of behavioural antagonists in the interspecific relations between Y.rorellus and sympatric closely related species are discussed.     

Pheromone-mediated mating disruption of Choristoneura rosaceana: is the most attractive blend really the most effective?

An attractive four-component pheromone blend containing a major component Z11-tetradecenyl acetate, and three minor components, E11-tetradecenyl acetate, Z11-tetradecenyl alcohol, and Z11-tetradecenyl aldehyde was tested as a mating disruptant against western Canadian populations of the obliquebanded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae), in organic apple orchards in British Columbia. Efficacy of this four-component blend was compared to that of partial pheromone blends containing the major component plus one or two minor components. A trapping experiment confirmed that, Conrel® fibre disruption dispensers containing the four-component blend were more attractive than disruption dispensers containing the two- or three-component partial blends. A small-plot protocol was followed to compare atmospheric treatments with these blends as mating disruptants at a release rate of 10 mg ha–1 h–1 and from 1000 dispensers ha–1. Mechanisms of mating disruption, such as false-trail following and camouflage of pheromone plumes, that may be evoked to a greater degree by an attractive blend, did not appear to augment the effectiveness of mechanisms invoked by the less attractive blends, as the proportion of mating among tethered females was equal in plots treated with these blends and was reduced by 85–90% compared to the nontreated control. When the four-component pheromone blend was tested at different release rates, mating disruption in small plots began to break down at a release rate of 1.3 mg ha–1 h–1 using a dispenser density of 1000 ha–1. Above 1.3 mg ha–1 h–1 there was no dose response in release rates tested and at release rates below this dose the proportion of tethered females mating was the same as in the nontreated control. The four-component pheromone blend was tested against, and found to be no more effective than, the two-component partial blend at the threshold release rate of 1.3 mg ha–1 h–1 when it was released from 1000 or 250 disruption dispensers. Our results suggest that disruption mechanisms evoked by the attractive blend did not enhance the mating disruption effect provided by the simple blend, therefore a two-component blend may be useful in an operational mating disruption program for C. rosaceana.