Circadian rhythmicity and development of the behavioural response to sex pheromone in male brown-banded cockroaches, Supella longipalpa

Abstract The effects of age, time of day and pheromone concentration on the responsiveness of the male brown-banded cockroach, Supella longipalpa (F.), to female volatile sex pheromone were examined. Male response increased with age and peaked 10 days after the imaginal moult. A diel periodicity of responsiveness was observed, with greater responses in the scotophase. The behavioural response was dose-dependent and its diel periodicity was most obvious when intermediate pheromone concentrations (e.g. 0.01 female equivalents) were used. Higher concentrations obscured the periodicity by eliciting greater responses in the photophase, while the lower concentrations did so by decreasing the response in the scotophase. Following entrainment to a LD 12:12 h cycle, the periodicity of response to 0.01 female equivalents of pheromone persisted for at least 54 h under continuous darkness, further demonstrating a true circadian rhythm.     

Temperature-mediated sex pheromone production and response of the American cockroach

Temperature affects the production of and response to sex pheromone by the American cockroach. Female sex-pheromone production is directly related to acclimation temperature and rapidly decreases 1 week after acclimation to lower temperatures. Production decreases slightly between weeks 1 and 2 and remains constant thereafter. Phermone-release rates are not affected by substrate temperature. Male response to a 1 ♀-equivalent hr pheromone dose is also temperature dependent. Maximal response was obtained at 26.7°C. Acclimated males were able to perceive pheromone (EAG technique) and capable of movement at lower temperatures (10°C), suggesting central nervous system inhibition.     

梨小食心虫性诱剂附加农药诱杀器的设计

通过田间对梨小食心虫雄蛾着陆行为观察,设计出4种诱杀器,其中双翼式和台式最适于雄蛾降落。诱杀器的材料以PVC塑板为好。涂在诱杀器上的缓释型菊酯农药,开始对雄蛾有较强的忌避作用,但在田间暴露10天后,雄蛾着陆率可在80％以上,让雄蛾接触在田间暴露的药膜5秒左右,开始雄蛾可在10分钟内被击倒,当药膜在田间暴露80天后,仍可击倒雄蛾,所以这种性诱剂加农药的诱杀器有可能用来防治梨小食心虫或其它小型害虫。     

Temperature modulation of behavioural thresholds controlling male moth sex pheromone response specificity

ABSTRACT. The response specificity of male Oriental fruit moths, Grapholita molesta (Busck) and pink bollworm moths, Pectinophora gossypiella (Saunders), to different blends and doses of pheromone is altered dramatically by temperature. When acclimated and tested in a flight tunnel at 20^oC males of both species exhibit a high degree of specificity, with peak response levels occurring to a narrow range of blend-dose combinations close to the natural blend. When tested at 26^oC, however, males exhibit a significantly lower degree of specificity, with peak response occurring to a broader range of treatments. The change in response specificity results from shifts in behavioural threshold effects influencing plume orientation and initiation of upwind flight, as well as from arrestment of upwind flight, occurring later in the flight sequence. The observed changes in male behaviour suggest that the effect of temperature is directly on neural pathways involved in the perception of odour, and not simply the result of an increase in motor activity or a significant change in the release rate of the pheromone. The results support the threshold hypothesis for pheromone perception (Roelofs, 1978) as a general principle in the Lepidoptera, but also show that the degree of response specificity can be significantly affected by temperature.     

Rate and diel periodicity of pheromone emission from female gypsy moths, (Lymantria dispar) determined with a glass-adsorption collection system

The rate of pheromone emission from wild and laboratory-reared gypsy moth (Lymantria dispar) (Lepidoptera: Lymantriidae) virgin females was determined with an all-glass aeration apparatus. This device incorporated a bed of 1-mm glass beads to extract entrained pheromone from the air flowing over the protruded gland. The temporal pattern of emission was established by monitoring individual females after eclosion for 24 consecutive 2-hr intervals.At a constant 24°C, both wild and laboratory females exhibited a similar diel periodicity of pheromone emission. The mean release rate increased after onset of photophase, generally attained maximal levels between 1600 and 2200 hr and declined during scotophase. Pheromone was released continuously and the mean daily emission increased with age for both wild and laboratory moths. The mean emission rate over the 48-hr monitoring interval was $\text{15.4 ng}\text{2 hr}$ for wild females vs $\text{14.7 ng}\text{2 hr}$ for laboratory moths. The peak emission from 2-day-old laboratory moths was ca.$\text{28 ng}\text{2 hr}$ compared with the ca.$\text{25 ng}\text{2 hr}$ released by their wild counterparts.The calling periodicity of laboratory females was determined at a constant 24°C and under a natural temperature rhythm. At 24°C, the proportion of females calling exceeded 45% throughout the diel period, whereas under the temperature rhythm, calling was virtually eliminated by temperatures below 15°C, indicating that temperature acts as an exogenous cue to modify the expression of the calling rhythm and thus potentially the periodicity of pheromone emission.     

Effects of temperature and light on calling in the tiger moth Holomelina lamae (Freeman) (Lepidoptera: Arctiidae)

ABSTRACT. In Holomelina lamae Freeman daily eclosion of adults is gated, with males emerging before females. By advancing the onset of photophase and by delaying the onset of scotophase, it was demonstrated that lights-on acts as the main phase-setting cue for calling. Few females call on the day they eclose. Calling is initiated c. 9 h after the onset of photophase in 2-day-old females, and shifts to earlier times in older females. The duration of calling also increases with age. That calling is controlled by an endogenous circadian clock is indicated by its persistence in continuous light (LL) and dark (DD). In LL calling is dampened rapidly, but a single scotophase re-entrains the rhythm. Decreases in temperature advance the onset of calling and the mean hour of calling, while increases in temperature delay both. However, the magnitudes of such phase-shifts depended upon hour of the photoperiod. Moreover, cooling and heating appears to exert both transient and long-term effects on the calling rhythm. An 8 h period at a reduced temperature in LL induces calling in females whose calling is dampened, and entrains the calling rhythm. Females maintained in DD from second instar larvae to the adult stage exhibit a circadian calling rhythm set by eclosion.     

楸螟行为观察及性信息素粗提物生物活性测定

本文报道了楸螟Omphisa plagialis Wileman的羽化昼夜规律和（25±1）℃、光周期L∶D=14∶10条件下的雌蛾求偶节律,用Y形管法测定了雄蛾对雌蛾性信息素粗提物的行为反应。结果表明：楸螟的羽化发生在10：00到23：00之间;雌蛾在羽化当天即开始求偶,求偶行为集中在暗周期的23：30到01：30之间。采用正己烷提取的雌蛾性信息素粗提物存在性信息素成分,对雄蛾具有一定的引诱活性。     

Environmental regulation of female calling and male pheromone response periodicities in the codling moth (Laspeyresia pomonella)

Codling moth female calling and male pheromone responsiveness under the defined conditions of 23°C and light:dark (LD) 16:8 occurred primarily during scotophase. Under either continuous photophase or scotophase females called with periodicities very similar to their periodicity under the LD cycle, indicating that the rhythmicity is circadian. Male response rhythmicity was maintained under continuous photophase. A decrease in the temperature from 23° to 16°C resulted in a reduction in the proportion calling when the decrease in temperature occurred during scotophase and a shift of maximal calling into photophase when the decrease in temperature occurred 3 hr prior to the initiation of scotophase. Decreases of temperature from 23° to 16°C and of light intensity did not produce similar shifts in the periodicity of male upwind orientation. Of 6 pheromone dosages from 10^?5 to 10² μg, 10^?1 and 10⁰ μg elicited the most male upwind orientation.     

An aphid circadian rhythm: Factors affecting the release of sex pheromone by oviparae of the greenbug, Schizaphis graminum

Adult oviparae of Schizaphis graminum emit a sex pheromone from scent plaques on the metathoracic tibiae, as do oviparae of other aphid species. By allowing males to walk along a wire walkway, a turning response and an increased rate of antennation were observed when the aphids were within 1–2 cm of a pheromone source. Adult males responded but 4th instar larval males did not.The onset of the release of the pheromone by the oviparae is triggered by the initiation of the light phase in a LD-cycle, and is governed by a circadian rhythm with a free-running period of 25.6 hr under continuous illumination of 15 lx. Daily pheromone release peaks 4–7 hr after lights-on and reaches a maximum on days 6–8 of adulthood.     

Neurological influences on pheromone release and calling behaviour in the gypsy moth, Lymantria dispar

ABSTRACT. Surgical removal of the brain or disconnection of the last abdominal ganglion from the ventral nerve cord prevented sex pheromone release in female Lymantria dispar (L.) (Lymantriidae), as assayed by the male wing-fanning response. The calling behaviour continued to occur in individuals whose terminal abdominal ganglion had been thus isolated, however, indicating that the neural mechanisms controlling calling function independently in the last abdominal ganglion.     

A reinvestigation of the role of (E)-2-hexenal in female calling behaviour of the polyphemus moth (Antheraea polyphemus)

Calling periodicities in the saturniid moth Antheraea polyphemus were similar in virgin females that were intact, intact and exposed to (E)-2-hexenal and antennectomised. Virgin females and antennectomised virgin females were of comparable attractiveness to males in the field.     

The effects of different blend ratios and temperature on the active space of the Oriental fruit moth sex pheromone

Abstract The wing-fanning activation response of male Oriental fruit moths (OFM), Grapholita molesta (Busck), in the field to the three-component pheromone containing the female-produced ratio of components (Z8-12:OAc + 6% E8-12:OAc + 3% Z8-12:OH) was compared with the response to blends containing 2,10 and 20% E with 3% OH, and the 6% E blend containing 30 and 100% OH. Comparisons were made over three temperature ranges: 15–17, 20–21 and 26–28^oC. Both the maximum response distance and male response specificity were significantly altered by changes in odour quality as well as temperature. For blends containing different Z/E ratios the maximum response distance increased significantly with temperature. Response specificity was most pronounced at the 20–21^oC range, with males displaying a lower threshold for the natural 6% E ratio, evidenced by the fact that fewer males responded and at closer distances to the source with off-ratios. At 26–28^oC response specificity for the Z/E ratios was much reduced, primarily due to more males activating to off-ratios. With blends containing different proportions of Z8-12:OH in the 6% E blend, increasing temperature increased the maximum response distance for all treatments, but in addition increasing the proportion of OH alone from 3% to 30% significantly increased the maximum response distance over the three temperature ranges tested. This increase occurred without affecting the proportion of responders or the distribution of response distances around the mean value. However, with 100% OH added to the blend, whereas male response was high at 20–21^oC, the distribution of response distances was significantly more variable than with 3% or 30%, and male response was eliminated or very low at 15–17^oC and 26–28^oC. Our results support previous studies showing that peak response levels in this species are dependent on male perception of the natural blend of components, and that males have a high degree of specificity for the qualitative properties of the pheromone. However, the present results also extend those of previous flight tunnel tests in which response specificity was most pronounced in the upwind flight phase of the sequence, by showing that male OFM also display a     

Control of Grapholita molesta (Busck, 1916) (Lepidoptera: Tortricidae) with entomopathogenic nematodes (Rhabditida: Heterorhabditidae,Steinernematidae) in peach orchards

Oriental fruit moth Grapholita molesta (Busck, 1916) (Lepidoptera: Tortricidae) is considered a major pest in temperate fruit trees, such as peach and apple. Entomopathogenic nematodes (EPNs) are regarded as viable for pest management control due to their efficiency against tortricid in these trees. The objective of this study was to evaluate the effectiveness of native EPNs from Rio Grande do Sul state against pre-pupae of G. molesta under laboratory and field conditions. In the laboratory, pre-pupae of G. molesta were placed in corrugated cardboard sheets inside glass tubes and exposed to 17 different EPNs strains at concentrations of 6, 12, 24, 48 and 60 IJs/cm² and maintained at 25 °C, 70 ± 10% RH and photophase of 16 h. Insect mortality was recorded 72 h after inoculation of EPNs. Steinernema rarum RS69 and Heterorhabditis bacteriophora RS33 were the most virulent strains and selected for field application (LC₉₅ of 70.5 and 53.8 IJs/cm², respectively). Both strains were highly efficient under field conditions when applied in aqueous suspension directed to larvae on peach tree trunk, causing mortality of 94 and 97.0%, respectively.     

Dual pheromone dispenser for combined control of codling moth Cydia pomonella L. and oriental fruit moth Grapholita molesta (Busck) (Lep., Tortricidae) in pears

Codling moth (CM) and oriental fruit moth (OFM) are very important orchard pests worldwide, and particularly in Victoria, Australia, where both species damage pome fruit. Individually CM and OFM have been controlled successfully by pheromone‐mediated mating disruption, but treatment of pome fruit with full registered rates of two individual hand‐applied dispensers for CM and OFM could be uneconomical for growers. Field trials conducted over three seasons in plots sprayed with insecticides consistently demonstrated that dual Isomate C/OFM TT dispensers, designed to disrupt both CM and OFM, were as effective as Isomate CTT and Isomate OFM Rosso dispensers applied individually in pears for control of CM and OFM respectively. The dual‐ and individual‐species dispensers reduced moth catches and fruit damage to a similar degree. The results suggest that combined control of CM and OFM in pears by applying dual Isomate C/OFM TT dispensers at the full‐recommended rate of 500 dispensers per hectare will be more economical than use of individual species dispensers, because the price and application cost of dual dispenser is about half that for individual dispensers.     

Effect of varying proportions of the alcohol component on sex pheromone blend discrimination in male Oriental fruit moths

ABSTRACT. The flight response of individual male Oriental fruit moths, Grapholitha molesta (Busck), was observed in a sustained-flight tunnel to 100 blend–dosage combinations of the three sex pheromone components: (Z)- and (E)-8-dodecenyl acetate and (Z)-8-dodecen-l-ol (1, 3, 10, 30 and 100 μg of Z8-12: AC with, 2, 6, 10, 20% E and, 0, 3, 10, 30 or100% OH alcohol added). Complete flights to the source were observed only to blend combinations containing all three components. Males exhibited highest response levels to two dosages (3 and 10μg) of the natural 6% E blend and these levels were relatively unaffected by changes in the proportion of Z8-12: OH. Certain treatments surrounding the peak area also elicited high response levels compared to the 6% E treatments, but these were strongly dependent on the proportion of OH in the blend. Hierarchical cluster analysis was utilized to compare and group treatments that elicited similar levels of response over the entire flight sequence and to derive an area of blend-dosage combinations within the matrix tested that elicited peak levels of attraction. Analysis of the response patterns for suboptimal treatments adjacent to the area of optimal attraction showed that response specificity was controlled by two major effects on flight behaviour, one occurring early in the flight sequence affecting male orientation to the odour plume, and the other occurring later in the sequence as an arrestment of upwind flight. These effects were strongly influenced by changes in the OH component and the E isomer, with the latter playing the critical role in effecting flight behaviour. Temporal analysis of the flight response to treatments within the optimal area showed that whereas the % OH did not significantly affect the magnitude of response, increasing the level of Z8-12: OH in the blend did significantly increase the duration of each phase of the behavioural sequence. Considering both the magnitude and temporal aspects of male response, optimal attraction in male OFM was best characterized by a small area of treatments around the peak 6% E blends that contained 10% OH. Finally, field tests showed a high degree of correlation between trends in response with changing proportion of Z8-12: OH as observed in the flight tunnel. Peak dosages were generally higher in the field, however, compared to those in the flight tunnel.     

The effects of mating,exogenous juvenile hormone and a juvenile hormone analogue on pheromone titre,calling and oviposition in the omnivorous leafroller moth (Platynota stultana)

Mating in Platynota stultana resulted in the termination of calling, the gradual reduction of pheromone in the pheromone glands to non-detectable levels (<0.1 ng/♀) within 14 h, and oviposition of the first batch of eggs 20–24 h after copulation. Decapitation of virgin females resulted in a similar decline in pheromone titre, and also eliminated oviposition and calling. Pheromone production appears to be controlled via the head. Mating probably terminates neural or hormonal input required for pheromone production and/or removes neural or hormonal inhibition of pheromone degradation. A juvenile hormone analogue (ZR-512) and juvenile hormones I, II and III applied exogenously to virgin females elicited oviposition comparable to mated females and terminated calling within 48 h. The juvenile hormone analogue also appeared to block pheromone production in virgin females. These results suggest that juvenile hormone may be involved in the switch from virgin to mated behaviour in this species.     

Sublethal effects of neuroactive compounds on pheromone response thresholds in male oriental fruit moths

The pheromone-mediated flight behavior of male Oriental fruit moths in a sustained-flight tunnel was observed after males were treated topically with sublethal concentrations of permethrin, carbaryl, chlordimeform, dieldrin, octopamine, serotonin, yohimbine, and cyproheptadine. With the exception of serotonin all compounds were found to disrupt one or more specific elements of the male precopulatory flight sequence. Among the insecticides, dieldrin was least active, whereas permethrin, carbaryl, and chlordimeform induced unique effects at specific phases of the sequence. Octopamine induced a hypersensitivity to the olfactory signal and mimicked one of the effects observed with chlordimeform. Yohimbine and cyproheptadine significantly decreased moth activation to the chemical signal but did not alter flight performance in responding moths. Yohimbine and cyproheptadine also reversed the effects induced by octopamine. The results of our study show that the complex precopulatory sequence of behaviors exhibited by males is very sensitive to sublethal concentrations of a range of neuroactive compounds.     

The alteration of calling behaviour by female Choristoneura fumiferana when exposed to synthetic sex pheromone

The calling behaviour of 0-day-old and 2-day-old virgin female eastern spruce budworm moths, Choristoneura fumiferana (Clem.) was observed during exposure to different concentrations of the synthetic female sex pheromone. Regardless of whether the females were exposed to pheromone or not, there were two daily peak periods of calling activity, one in the afternoon and the other after dusk. Those moths exposed to pheromone commenced calling at least 2 h earlier during their first peak of calling activity. Also, a greater number of moths engaged in calling when exposed to pheromone than those under control conditions.

---

Résumé Des expériences ont été menées au laboratoire pour examiner si la phéromone sexuelle synthétique de C. fumiferana pouvait altérer le comportement d'appel de femelles vierges de la même espèce. Un appareil a été conçu pour libérer alternativement — à des intervalles déterminés — un courant d'air portant de la phéromone sexuelle ou un courant d'air pur, dans plusieurs flacons contenant des femelles vierges. Le comportement sexuel de mouches de 0 et 2 jours a ainsi été observé. Indépendamment de l'exposition des femelles à la phéromone, il y avait deux pics quotidiens d'activité d'appel, l'un dans l'après-midi et l'autre après le crépuscule. Des papillons exposés au 1×10^–6 mg ou 1×10^–3 mg de phéromone avaient un premier pic d'appel au moins antérieur de 2 heures par rapport aux témoins. Ainsiily a vait un plus grand nombre de papillons engagés dans une activité d'appel en présence de la phéromone que chez les témoins.

     

蛾类昆虫雄性信息素及其功能

昆虫性信息素是两性通讯系统的基础,其中雄性信息素的研究相对较少。本文综述了蛾类昆虫雄性信息素的研究进展。迄今已鉴定出40余种蛾类昆虫的雄性信息素,其行为学功能主要有对雌性的引诱和激欲、对同种雄性的抑制及种间隔离等。     

Manoeuvres used by flying male oriental fruit moths to relocate a sex pheromone plume in an experimentally shifted wind-field

ABSTRACT. In a wind-field experimentally shifted in direction by 35d?, flying male Grapholita molesta (Busck) zigzagging upwind either maintained contact with a pheromone plume and followed it across during the shift or lost it and commenced casting at c. 90d? across the shifting windline to locate it eventually in its new position. Males accomplished both of these results by integrating the previously described systems of optomotor anemotaxis and self-steered counterturning, but with faster reaction-times to pheromone on and off than heretofore calculated for this species. We found no evidence that males following the plume across used chemotaxis as proposed for another species, Rather, the sawtoothed-shaped tracks were a result of the anemotactic and counterturning systems responding rapidly and reiteratively to each loss and gain of pheromone along the plume in the shifting wind. The response to an increase or decrease in pheromone concentration by males was to change their course angle to more upwind or more crosswind, respectively, on the very first reversal (within c. 0.15 s) after the concentration changed. Because males adjusted their airspeeds more slowly to changes in concentration, the groundspeeds along the more upwind-orientated legs were lower than those along cross-wind legs, contributing to the sawtoothed shape of tracks of plume-followers. The self-steered counterturning programme also reacted quickly to concentration changes, the reversal intervals tending to be shorter following each contact with pheromone than after each excursion into cleaner wind. Following casting after losing the plume, males relocating the pheromone plume exhibited an upwind ‘surge’ of narrow zigzagging flight because on the first leg in the plume they steered a course more directly upwind than on the previous leg and increased the frequency of counterturning to its highest value while maintaining the relatively high airspeed acquired while casting.