Interactions between Acetoin,a Plant Volatile,and Pheromone In <Emphasis Type="Italic">Rhynchophorus palmarum</Emphasis>: Behavioral and Olfactory Neuron Responses

Aggregation of Rhynchophorus palmarum weevils on host plants is mediated by a male pheromone (rhynchophorol: R) and host-plant volatiles (PVs) acting in synergy. Synthetic PV blends synergizing pheromone contain acetoin (A) and ethyl acetate (EtAc). R, A, and EtAc are detected by specialized olfactory receptor neurons (ORNs). In addition, particular types of ORNs are tuned to both A and R. To specify the role played by acetoin in pheromone perception, we recorded the responses of ORNs to 100 ng of A or R presented either separately or mixed. Behavioral responses to R, A, and EtAc were studied in a four-armed olfactometer and by field trapping. We screened 59 R-, A-, and AR-tuned ORNs by recording specific responses to odors presented either separately or mixed. Stimulations by blends elicited complex response profiles from the three ORN types: some gave synergistic responses, others were inhibited, and the remainder responded as though both odors were detected independently. Several gave either a weak or no response to a first stimulation by R, but responded clearly to a second stimulation after an intercalary stimulation by A. In the olfactometer, both sexes were more attracted to a blend of A + R (1 + 0.01 ng/sec) than to pure compounds, whereas EtAc did not enhance response to R. Pheromone-baited traps (1 mg/day) containing PV blends (650 mg/day) based on an ethanol/EtAc blend (1:1), plus either 5 or 10% A, or a more complex reference blend, or sugarcane (natural pheromone synergist), caught similar numbers of weevils and about twice as many insects as a control ethanol/EtAc blend. Traps with only pheromone caught about 10 times fewer insects. Behavioral results support the role of acetoin as a pheromone synergist for R. palmarum, and electrophysiological data provide evidence of modulation of peripheral sensory responses to pheromone by acetoin. Sexual dimorphism was observed neither at the ORN nor at the behavioral levels.     

Evolution of behavioral responses to sex pheromone in mutant laboratory colonies ofTrichoplusia ni

Male cabbage looper moths,Trichoplusia ni, from two colonies in which all females express an abnormal sex pheromone production phenotype were evaluated in a laboratory wind tunnel for upwind flight responses to the normal and abnormal sex pheromones. The abnormal sex pheromone blend consisted of 20 times as much (Z)-9-tetradecenyl acetate and 30-fold less (Z)-5-dodecenyl acetate compared to the normal pheromone blend. Initially, these males exhibited poor behavioral responses to the abnormal sex pheromone and maximum responses to the normal pheromone blend, indicating that there was no linkage between signal production and response. After 49 generations of laboratory rearing, males from the mutant colonies maintained good responses to the normal pheromone and increased their behavioral response to the abnormal sex pheromone to the same levels as for the normal pheromone. Over the same period, normal males maintained their preference for the normal pheromone. These results indicated that evolution had occurred in mutant colonies in favor of greater male responsiveness to the abnormal sex pheromone, resulting in the broadening of the response spectrum to pheromone blend ratios. This evolution presumably resulted from a mating advantage to those males that did not discriminate against mutant-type females in the mutant colonies.     

Male moth sensitivity to multicomponent pheromones: Critical role of female-released blend in determining the functional role of components and active space of the pheromone

In the present study male redbanded leafroller (Argyrotaenia velutinana), cabbage looper (Trichoplusia ni), and Oriental fruit moths, (Grapholita molesta), were tested in a flight tunnel to (1) the major pheromone component, (2) theZ/E pheromone component mixtures for Oriental fruit moth and redbanded leafroller, (3) and the female-released blends, over a series of dosages. Experiments were designed to test the hypothesis that male response downwind of a female is initiated by the major component and that minor components function only to elicit behaviors close to the female during close-range approach and courtship. The results did not support this hypothesis, but rather showed that males initiated upwind flight in significantly higher percentages to the complete blends of components, at all dosages, compared to single components or partial blends. Addition of minor components also signficantly enhanced male perception of the major component at lower dosages, resulting in completed flights to dosages of the major component that alone did not elicit any upwind flight. Our results support the concept that minor components function to enhance male sensitivity to the pheromone, and the specificity of the signal. Our results also support the hypothesis that the active space of the pheromone is a function of the upper and lower concentration thresholds for the blend of components, and not simply for the major component.     

Activity of perfluorobutyl-containing components in pheromone blend of cabbage looper moth,Trichoplusia ni

Substitution of nine hydrogens by nine fluorines in the butyl terminus of (Z)-7-dodecenyl acetate, the major component of the pheromone blend of the cabbage looper,Trichoplusia ni (Lepidoptera: Noctuidae), produces a highly volatile analog that is significantly less active as a replacement for the natural pheromone component, either alone or substituted in the sixcomponent pheromone blend of this species. When 9,9,10,10,11,11,12,12, 12-nonafluoro-(Z)-7-dodecenyl acetate (nf-Z7-12: OAc) was tested in a wind tunnel as a single component or as part of the six-component blend, males exhibited preflight activation, but they did not orient in the pheromone plume and fly upwind over a five-decade range of doses at levels comparable with those of the natural component and blend. Addition of large amounts of nf-Z7-12:OAc to the optimal dose of the natural blend also did not effect flight arrestment as when increased amounts of Z7-12:OAc were added. The nonafluoro (nf) analog of (Z)-9-tetradecenyl acetate, a minor component in theT. ni blend, did not affect male behavior when substituted in the sixcomponent blend, and substitution of both nf analogs in the natural blend produced the same results as observed for blends containing nf-Z7-12:OAc. The lack of an effect with substitution of nf-Z9-14:OAc is discussed in relation to the reported redundancy in the action of minor components inT. ni.     

Pheromone-mediated responses of male cabbage looper moths,Trichoplusia ni,following various exposures to sex pheromone or (Z)-7-dodecenol

Prolonged preexposure (three days) of maleTrichoplusia ni to its six-component sex pheromone blend or its major pheromone component, (Z)-7-dodecenyl acetate, reduced subsequent upwind flight responses to a pheromone source. Preexposure to (Z)-7-dodecenol increased upwind flight responses to a pheromone source combined with (Z)-7-dodecenol. The impact of long-term preexposures was moderate when compared to the more immediate effects of background noise. When (Z)-7-dodecenyl acetate was presented as background noise, all maleT. ni failed to respond to a plume of the full pheromone blend. However, most moths succeeded in locking on to the pheromone plume and contacting the pheromone source in the presence of the five minor pheromone components as background noise. When (Z)-7-dodecenol was released as background noise the response rate to a pheromone source containing (Z)-7-dodecenol was increased dramatically. This indicates that males became adapted to (Z)-7-dodecenol while responding to the pheromone source. The results of this study indicate that both long-term preexposure treatments and immediate exposure to background noise can limit the ability of maleT. ni to respond to sex pheromone sources.     

Trade-Off Between Sensitivity and Specificity in the Cabbage Looper Moth Response to Sex Pheromone

The evolution of male moth responses to pheromone blends may be constrained by a trade-off between two response traits: sensitivity and breadth of response. Population genetic simulations predict that if sensitivity and breadth of response are negatively correlated (i.e., a trade-off exists), then selection will favor males with narrow response phenotypes and high sensitivity. Although sensitivity-breadth of response trade-offs are generally assumed to exist and are implicit to the shape of male preference function, this study is the first to provide empirical support measuring behavior at the level of the individual. Previous studies with the cabbage looper, Trichoplusia ni, have documented the existence of a mutant pheromone strain. While mutant females produce a pheromone blend significantly different from wild-type females, mutant males respond equally to the wild-type and mutant pheromone blends. This study used wind tunnel bioassays to document that relative to wild-type males, mutant males had broader response profiles but lower pheromone sensitivity. While wild-type male responses were highest to the wild-type pheromone blend, mutant males did not discriminate among pheromone blends. These results are consistent with a trade-off between breadth of response and sensitivity. Pure wild-type and mutant lines were crossed and hybrid males assayed. Both hybrid types (maternal wild-type and maternal mutant hybrids) responded similarly. Hybrid males had response profiles similar to wild-type males and the reduced sensitivity observed in mutant males. These results suggest a possible hybrid disadvantage and a putative mechanism for reinforcement of male pheromone response traits.     

Flight Tunnel Response of Male European Corn Borer Moths to Cross-Specific Mixtures of European and Asian Corn Borer Sex Pheromones: Evidence Supporting a Critical Stage in Evolution of a New Communication System

Previous flight tunnel studies showed that 3–5 % of male European corn borer (ECB) moths, Ostrinia nubilalis, could fly upwind and make contact with sources releasing the sex pheromone of the closely related Asian corn borer (ACB), Ostrina furnacalis, [2:1 (Z)-12-tetradecenyl acetate (Z12-14:OAc) : (E)-12-teradecenyl acetate (E12-14:OAc)] and that 2–4 % of ACB males could similarly fly upwind to the sex pheromone blends of the ECB Z- [97:3 (Z)-tetradecenyl acetate (Z11-14:OAc) : (E)-tetradecenyl acetate (E11-14:Ac)] and E-strains (1:99 Z/E11-14:OAc) pheromones. The results supported the hypothesis that the evolution of the ACB pheromone system from an ECB-like ancestor included a stage in which males could be attracted to the unusual females emitting Z12- and E12-14:OAc while retaining their responsiveness to the ancestral pheromone blend of Z11- and E11-14:OAc. Here, we showed further that ECB E-strain males exhibited upwind oriented flight and source contacts to sources containing all combinations of ECB and ACB components. Maximal response levels were observed with the E-strain 99:1 E11/Z11-14:OAc blend, and high response levels also were observed with two other blends containing E11-14:OAc as the major component (E11:E12 and E11:Z12). Upwind flight and source contact also occurred at lower levels with the remaining blend combinations in which Z11-, E12-, or Z12-14:OAc was the major component. Our current results support the hypothesis concerning the evolution of ACB from an ECB-like ancester by showing that males were able to respond to females producing either the 12–14:Ac isomers, 11–14:Ac isomers, or even mixtures of all four components     

Mating Disruption of Guatemalan Potato Moth Tecia Solanivora by Attractive and Non-Attractive Pheromone Blends

The behavioral mechanisms of mating disruption in Guatemalan potato moth Tecia solanivora were studied using the sex pheromone components, (E)-3-dodecenyl acetate, (Z)-3-dodecenyl acetate, and dodecyl acetate, formulated in a 100:1:20-ratio mimicking the female-produced blend, and in a 100:56:100 off-blend ratio. The mode of action of these two blends was tested in mating disruption experiments in the field and in a greenhouse, as well as in a laboratory wind tunnel. Field treatments with both blends at 80 g pheromone per ha reduced male attraction to trap lures baited with 100 μg of female sex pheromone. In mesh-house treatments, these two blends were equally effective at reducing male attraction to traps baited with live females and mating of caged females. Subsequent flight tunnel tests corroborated that both blends reduced attraction of naive males to calling females, and pre-exposure of males with either dispenser blend for 24 hr resulted in a strongly reduced response to calling females. The pre-exposure effect was reversible, with males again responsive after 24 hr in clean air. The two dispenser formulations produced a similar effect on male behavior, despite the differences in blend composition. One mating disruption dispenser formulated with either the female-blend or off-blend elicited the same rate of male upwind attraction in a wind-tunnel bioassay. Sensory overload and camouflage, therefore, are contributing mechanisms to mating disruption using either blend. The off-blend, which is more economical to synthesize, is a valuable tool for further development of mating disruption against this major pest of potatoes in Latin America.     

Field Attractants for <Emphasis Type="Italic">Pachnoda interrupta</Emphasis> Selected by Means of GC-EAD and Single Sensillum Screening

The sorghum chafer, Pachnoda interrupta Olivier (Coleoptera: Scarabaeidae: Cetoniinae), is a key pest on sorghum, Sorghum bicolor (L.) Moench (Poaceae), in Ethiopia. At present there is a lack of efficient control methods. Trapping shows promise for reduction of the pest population, but would benefit from the development of attractive lures. To find attractants that could be used for control of P. interrupta, either by mass trapping or by monitoring as part of integrated pest management, we screened headspace collections of sorghum and the highly attractive weed Abutilon figarianum Webb (Malvaceae) for antennal activity using gas chromatograph-coupled electroantennographic detection (GC-EAD). Compounds active in GC-EAD were identified by combined gas chromatography and mass spectrometry (GC-MS). Field trapping suggested that attraction is governed by a few influential compounds, rather than specific odor blends. Synthetic sorghum and abutilon odor blends were attractive, but neither blend outperformed the previously tested attractants eugenol and methyl salicylate, of which the latter also was part of the abutilon blend. The strong influence of single compounds led us to search for novel attractive compounds, and to investigate the role of individual olfactory receptor neurons (ORNs) in the perception of kairomones. We screened the response characteristics of ORNs to 82 putative kairomones in single sensillum recordings (SSR), and found a number of key ligand candidates for specific classes of ORNs. Out of these key ligand candidates, six previously untested compounds were selected for field trapping trials: anethole, benzaldehyde, racemic 2,3-butanediol, isoamyl alcohol, methyl benzoate and methyl octanoate. The compounds were selected on the basis that they activated different classes of ORNs, thus allowing us to test potential kairomones that activate large non-overlapping populations of the peripheral olfactory system, while avoiding redundant multiple activations of the same ORN type. Field trapping results revealed that racemic 2,3-butanediol is a powerful novel attractant for P. interrupta.     

More Rare Males in <Emphasis Type="Italic">Ostrinia</Emphasis>: Response of Asian Corn Borer Moths to the Sex Pheromone of the European Corn Borer

A previous flight tunnel study showed that 3–5% of European corn borer (ECB) moths, Ostrinia nubilalis (Z/E11-14:OAc), could fly upwind and make contact with sources releasing the sex pheromone of the related Asian corn borer (ACB), Ostrinia furnacalis (2:1 Z/E12-14:OAc). In this study, we show that rare males (3–4%) are also present in South Korean ACB that respond to the sex pheromone blends of the ECB UZ (97:3 Z/E11-14:OAc) and BE (1:99 Z/E11-14:OAc) pheromone races. We also show that the upwind flight response of a significant proportion of male ACB was antagonized by the addition of 1% Z9-14:OAc to the ACB blend, a compound that also antagonizes the upwind flight of ECB males. Male ACB flight behavior was not, however, affected by adding either of the ECB blends to the ACB blend, or by the addition of 50% 14:OAc, a compound identified from female pheromone glands of ACB and a number of other Ostrinia species. Additional flight tunnel tests with ACB to study the comparative aspects of ECB and ACB pheromone response specificity showed that male ACB exhibited maximal levels of upwind flight and source contact with doses of pheromone (30 and 100 μg on rubber septum sources) that also elicited maximal levels in the two ECB pheromone races. The maximal level of source contact for ACB (66%) was lower than observed with the UZ race of ECB to its pheromone blend (>95%), but comparable to those for the BE race of ECB (65–70%). Male ACB also flew upwind in high proportions to a broader range of ratios of Z/E12-14:OAc (80:20 to 20:80) than was previously observed for either of the ECB races.     

Specificity of male response to multicomponent pheromones in noctuid mothsTrichoplusia ni andPseudoplusia includens

The response of male cabbage looper (CL) and soybean iooper (SBL) moths was observed in the flight tunnel and measured in field tests to the six-component CL pheromone, the five-component SBL pheromone, and toZ7–12: OAc, the major component common to each pheromone. In both the flight tunnel and the field, male CL exhibited significantly greater levels of response to their six-component blend than toZ7–12: OAc alone. A low level of cross-attraction of male CL to the SBL pheromone was observed in both the flight tunnel and the field, but it was quantitatively and qualitatively similar to their response toZ7–12: OAc alone. Thus the minor components of the SBL blend did not appear to disrupt the flight behavior of male CL. With respect to SBL, in the flight tunnel males also exhibited a greater level of response to the five-component blend compared toZ7–12: OAc, but in the field their response was not significantly different to either treatment. There was also a low level of cross-attraction of male SBL to the CL blend, but this appeared to involve a significant arrestment effect on the upwind flight of males, as well as a difference in male sensitivity to the blend of components compared withZ7–12: OAc alone. The observed arrestment effect may have been due to male perception of one or more minor components of the CL pheromone. The results show that the multicomponent pheromones of these species function effectively as specific mating signals and that discrimination of odor quality by male moths can occur as the result of minor components affecting male sensitivity or their upwind flight response to the pheromone.     

Effects of host plant,Gossypium hirsutum L., on sexual attraction of cabbage looper moths,Trichoplusia ni (Hübner) (Lepidoptera: Noctuidae)

Unmated female or male cabbage looper moths,Trichoplusia ni (Hübner), were attracted more often in a flight tunnel to a cage with moths of the opposite sex and a bouquet of cotton foliage. Increased sexual attractiveness of females with plants may be a result of stimulation of pheromone release in response to plant odor, since more males were attracted when odor of cotton foliage was passed over females than when odor of females was passed over cotton foliage before venting into the flight tunnel. Increased sexual attractiveness of males with plants is due in part to host odor enhancement of female attraction to male pheromone, since more females were attracted to synthetic male pheromone (a blend of enantiomers of linalool and isomers of cresol) and a cotton leaf extract than were attracted to male pheromone alone. A short synthesis procedure was developed for (S)-(+)-linalool, the major component of the male sex pheromone, isolated from hair pencils, used in these tests.     

Role of sex pheromone components in behavioral reproductive isolation betweenAutographa gamma (L.) and eitherTrichoplusia ni (Hübner) ORChrysodeixis chalcites (Esp.) (Lepidoptera: Noctuidae: Plusiinae)

The composition of theAutographa gamma sex pheromone was reexamined and only (Z)-7-dodecenyl acetate and (Z)-7-dodecenol were identified by capillary GC, GC-MS, and dimethyl disulfide derivatization and subsequent GC-MS analysis. The fatty acid content of the pheromone glands was also studied, and a series of saturated and unsaturated acids was identified. However, most of the related pheromonal compounds were not detected. The male response to the pheromone components was studied in a flight tunnel and compared with the response to calling females. The best synthetic baits evoked a response similar to that observed to the virgin females, but males spent significantly more time at calling females than at the synthetic baits. The preferred synthetic baits consisted of (Z)-7-dodecenyl acetate alone or of a blend with 5% (Z)-7-dodecenol. Increasing the relative amount of the alcohol caused a gradual reduction in male response, particularly in the last steps of the courtship sequence. The addition of the minor sex pheromone components of the sympatric Plusiinae species,Trichoplusia ni andChrysodeixis chalcites, to theA. gamma pheromone was also investigated in the flight tunnel. Some of these components exhibited a significantly antagonistic effect on theA. gamma male courtship behavior. The most potent antagonists were (Z)-5-dodecenyl acetate and (Z)-9-tetradecenyl acetate. The response ofA. gamma andT. ni males to conspecific and heterospecific females was also compared in the flight tunnel. WhereasA. gamma males were attracted only to their conspecific females, a small percentage ofT. ni males were also attracted toA. gamma females and 11% performed the whole courtship sequence.Contribution from the Agricultural Research Organization (ARO), No. 3459-E, 1992 series.     

Filamentous nature of pheromone plumes protects integrity of signal from background chemical noise in cabbage looper moth,Trichoplusia ni

(Z)-7-Dodecenol failed to interrupt pheromone-mediated anemotactic responses by male cabbage looper moths,Trichoplusia ni (Hübner) (Lepidoptera: Noctuidae) in a wind tunnel when released 5 cm crosswind on both sides of the pheromone source or 10 cm upwind of the source to create an overlapping plume downwind. Significant inhibitory effects of (Z)-7-dodecenol were observed when released with the six-component pheromone blend from the same septum or abutting septa. These results indicate that (Z)-7-dodecenol needs to be received simultaneously with the pheromone blend to inhibit the anemotactic responses of males to the sex pheromone. We suggest that this feature and the filamentous nature of pheromone plumes render pheromone signals relatively protected from background chemical noise that may originate from pheromone plumes of other insect species. Unless filaments from a pheromone signal and an inhibitor arrive simultaneously, the integrity of the signal is maintained.     

Peripheral coding of sex pheromone and a behavioral antagonist in the Japanese beetle,Popillia japonica

Male antennae of the Japanese beetle, Popillia japonica, possess olfactory receptor neurons ORNs cocompartmentalized in the same sensilla placodea, one tuned to the sex pheromone, (R)-japonilure, and the other to the detection of a behavioral antagonist, (S)-japonilure. In-depth electrophysiological experiments revealed mutual inhibitory and synergistic effects in ORNs stimulated simultaneously with the two semiochemicals. The olfactory system of P. japonica exhibited a remarkable ability to discriminate completely coincident strands of pheromone and behavioral antagonist from strands of the two semiochemicals temporally isolated (by 1.5–3 msec). The mutual inhibition was reflected mainly by the delay of onset or total lack of spikes and by the significant increase in the rise time of potentials generated by blends of (R)- and (S)-japonilure. In contrast, synergist ORNs showed no neural activity (spikes) when stimulated with either the sex pheromone or the behavioral antagonist, but showed clear responses to blends of the two semiochemicals. Evidence for mixture-suppressed responses was observed not only in the Japanese beetle, but also in the Osaka beetle, Anomala osakana, and the Oriental beetle, Exomala orientalis, thus suggesting that it is a common feature in the sensory physiology of scarab beetles.     

Interpopulational variation in emitted pheromone blend of cabbage looper moth,Trichoplusia ni

Female cabbage looper moths,Trichoplusia ni, from laboratory colonies initiated from three locations across the United States emitted similar quantities and blend ratios of the six known pheromone components. In contrast, females from a long-established laboratory colony emitted a greater proportion of four of the five minor components relative to the major component, (Z)-7-dodecenyl acetate; only the relative proportion of 11-dodecenyl acetate was similar in all of the populations sampled. Females from this population emitted (Z)-7-dodecenyl acetate at a rate similar to that from females from field-collected colonies. Within each population there were highly significant correlations among the quantities of pheromone components of similar molecular weights. Correlations between components of different molecular weights were not as great, but often were significant. Similarities of blend ratios among field populations may indicate that the chemical signal in this species is conservative. The difference of the blend ratios in our laboratory population from the other populations may indicate a decrease in the intensity of selection pressure that usually would maintain these values.     

Identification of floral compounds fromAbelia grandiflora that stimulate upwind flight in cabbage looper moths

Four major volatile components emitted from flowers ofAbelia grandiflora were identified based on retention time using two capillary columns of different polarities and electron impact mass spectrometry. These are phenylacetaldehyde, benzaldehyde, 2-phenylethanol, and benzyl alcohol. A blend of these compounds was as effective as a cluster of flowers in stimulating upwind flight by maleTrichoplusia ni to the source in a wind-tunnel test. Phenylacetaldehyde or 2-phenylethanol were each as effective as the complete blend in stimulating source location by male moths. Attraction to a source of the synthetic blend was demonstrated in virgin males and females and mated males and females, but virgin moths of both sexes were more likely than mated moths to complete the sequence of behavioral responses necessary to locate the odor source.     

Heritable variation in pheromone response of the pink bollworm,Pectinophora gossypiella (Lepidoptera: Gelechiidae)

Heritability of variation in male pheromone response by pink bollworm moths,Pectinophora gossypiella (Saunders), was examined using a still-air, wing-fanning bioassay. Heritability (±SE) of overall responsiveness, as measured by the mean duration of wing fanning to the blend of pheromone components produced by females [4456 ratio of (Z, E)- to (Z, Z)-7,11-hexadecadienyl acetate], was 0.385 ± 0.095. Heritabilities of wingfanning duration to blends with 25 and 65%Z, E isomer were 0.377 ± 0.113 and –0.145 ± 0.103, respectively. These findings indicate an asymmetry in the genetic component of variation in response to pheromone blends with high and low proportions of theZ, E isomer. An index of response specificity for individual males was developed based on the response to an off-blend (either 25 or 65%Z, E isomer) relative to the response to the 44%Z, E blend. Heritabilities of response specificity were 0.117 ± 0.059 and –0.043 ± 0.067 for the 25 and 65%Z, E blends, respectively.     

Balanced Olfactory Antagonism as a Concept for Understanding Evolutionary Shifts in Moth Sex Pheromone Blends

In the sex pheromone communication systems of moths, both heterospecific sex pheromone components and individual conspecific pheromone components may act as behavioral antagonists when they are emitted at excessive rates and ratios. In such cases, the resulting blend composition does not comprise the sex pheromone of a given species. That is, unless these compounds are emitted at optimal rates and ratios with other compounds, they act as behavioral antagonists. Thus, the array of blend compositions that are attractive to males is centered around the characterized female-produced sex pheromone blend of a species. I suggest here that the resulting optimal attraction of males to a sex pheromone is the result of olfactory antagonistic balance, compared to the would-be olfactory antagonistic imbalance imparted by behaviorally active compounds when they are emitted individually or in other off-ratio blends. Such balanced olfactory antagonism might be produced in any number of ways in olfactory pathways, one of which would be mutual, gamma-aminobutyric-acid-related disinhibition by local interneurons in neighboring glomeruli that receive excitatory inputs from pheromone-stimulated olfactory receptor neurons. Such mutual disinhibition would facilitate greater excitatory transmission to higher centers by projection interneurons arborizing in those glomeruli. I propose that in studies of moth sex pheromone olfaction, we should no longer artificially compartmentalize the olfactory effects of heterospecific behavioral antagonists into a special category distinct from olfaction involving conspecific sex pheromone components. Indeed, continuing to impose such a delineation among these compounds may retard advances in understanding how moth olfactory systems can evolve to allow males to exhibit correct behavioral responses (that is, attraction) to novel sex-pheromone-related compositions emitted by females.     

Genetic aspects of interpopulational differences in pheromone blend of cabbage looper moth,Trichoplusia ni

The genetic basis of interpopulational differences in the pheromone blend emitted by the cabbage looper moth,Trichoplusia ni (Hübner), was examined by crossing individuals from a field-derived population (P₁) with individuals from a long-maintained laboratory colony (P₂). These colonies differed in the emission rate and relative proportions of four of the five known minor pheromone components, but not in the emission rate of the major component, (Z)-7-dodecenyl acetate (Z7-12Ac). These differences in pheromone blend were quantitatively small but biologically significant, because in the field, males responded preferentially to traps baited with a pheromone blend that is similar to that emitted by P₁ females relative to a blend similar to that emitted by P₂ females. In initial crosses, variation in the quantity and quality of pheromone blends among families of P₁, P₂, and F₁ hybrid females was examined. In F₁ females the relative proportions (quantity relative to the major component) of (Z)-5-dodecenyl acetate (Z5-12Ac) and (Z)-7-tetradecenyl acetate (Z7-14Ac) were intermediate to parental lines. In a second more extensive set of crosses, analyses included P₁, P₂, F₁, F₂, and selected backcrosses. The relative proportion of Z5-12Ac, Z7-14Ac, and Z9-14Ac emitted by F₁ females were intermediate to parental lines. The frequency distributions of relative proportions of these components emitted by females were not consistent with those expected under a single autosomal or sex-linked gene hypothesis, suggesting that more than one gene is involved in the quantitative differences in the pheromone blend.