Hormone-pheromone relationships of male Tenebrio molitor

Males of the beetle Tenebrio molitor produce a volatile sex pheromone which attracts females of the same species. The pheromone level peaks 8 days after emergence and then reaches a plateau. Elimination of endocrine centers by decapitating male adults 24 hr following adult ecdysis did not impair pheromone production. Treatment of decapitated males with juvenile hormone analogues did not make any detectable difference in the levels of pheromone activity. However, undecapitated males treated with juvenile hormone analogue showed a significant increase in pheromone activity when compared with those that had been decapitated and subsequently treated with juvenile hormone analogue. This observation is discussed in the light of published research on the effect of juvenile hormone on pheromone activity of females of T. molitor.     

OLFACTORY RESPONSES OF COTTON BOLLWORM,HELICOVERPA ARMIGERA TO SEX PHEROMONE AND PLANT VOLATILE COMPONENTS*

Abstract Electroantennograms (EAGs) were recorded from unmated, laboratory-reared male and female Helicover pa armigera (Lepidoptera:Noctuidae) adults to two sex pheromone components and a range of plant volatile components, some of which are known volatiles from leaves and fruits. The female-produced sex pheromone was not detected by other female moths. The sex pheromone components to which male moths responded previously show to possess biological activity. As for plant volatile components, a large degree of EAG response uniformity between male and female moth was observed. The greatest EAG responses of all plant volatiles tested were elicited by monoenic C-6 alcohol and aldehyde. They are constituents of the “general green-leaf odor” that emanates from most plants. The potential adaptive benefit of selective perception to sex pheromone and green-leaf volatile components is discussed.     

Isolation of sex pheromone mimic of the American cockroach by monitoring with male/female ratio in electroantennogram

The electroantennogram (EAG) technique was applied as a monitoring tool to isolate germacrene-D, a sex pheromone mimic of the American cockroach. Large values of the male/female ratio, and its index derived from EAG responses of adult males and females to a standard, indicated significant amounts of germacrene-D in plant fractions. Monitoring with behavioural assays and gas chromatographic analysis during the isolation confirmed that the EAG technique using the male/female index was a good indicator in the isolation of sexually active components.     

PHEROMONES AND CHEMICAL ECOLOGY OF LOCUSTS

Modern studies of chemical ecology and behaviour of the locusts Schistocerca gregaria and Locusta migratoria in the laboratory need to be more closely coupled with field experiments and observations. The life history relating to oviposition, transformation to gregarious phases, and adult maturation mediated by pheromones is reviewed. The principles of pheromone isolation and identification are discussed. The long-term effects of the gregarization pheromone on the physiology are presented, with discussion of morphological changes, chiasma frequency increases, and synchronization of moulting induced by the pheromone. Isolation of the purported gregarization pheromone, locustol, from faeces is discussed in regard to inconsistent effects. Other more immediate effects of the pheromone on the social (gregarious) behaviour and the isolation of possible pheromone components different from but related to locustol are presented. It is stressed that more rigorous isolation studies should be undertaken to resolve conflicting reports and methodological problems. The possibility of an anti-gregarization pheromone or solitarizing pheromone is discounted. The source and biosynthesis of locustol (or gregarization pheromone) from degradation of lignin by symbiotic bacteria is discussed. Theories of reception of the gregarization pheromone such as inhalation through the spiracles or sensory perception by the antennae are presented. Also an internal mechanism involving cAMP and/or corpora allata may be induced by gregarization pheromone to effect the physiological phase changes. The advantages to an individual of reception of the gregarization pheromone from a group of gregarious and pre-migrating locusts is discussed. Also the possible benefits of gregarious behaviour, phase polymorphism and migration are dealt with. An adult (sexual) maturation pheromone has long-term effects on reducing the period of maturation, and immediate effects on the behavioural vibration response. The epidermal source of the pheromone and glandular cells responsible for the production of the pheromone are discussed. The reception and internal mechanisms of response via the corpora allata are mentioned. The benefits to individuals of synchronized and rapid adult maturation in a gregarious group are considered. An oviposition-stimulating pheromone produced by the male accessory reproductive glands appears to be a proteinaceous substance of large molecular weight. On the other hand, an oviposition-aggregating pheromone volatilizes from epidermal areas of either sex and causes higher oviposition rates in the area of release. The behavioural and ecological aspects of this pheromone are discussed. Several other possible pheromones and semiochemicals are discussed, such as a long-range sex pheromone, sex-recognition pheromone, grass odours and feeding stimulants and deterrents. Several possible control strategies using locust pheromones are considered. The general conclusion is that the chemical isolation of the various pheromones is necessary before further progress can be achieved on the source and biosynthesis of pheromone, reception of pheromone, behavioural effects of pheromone, and control measures.     

Antennal responses of Cydia pomonella (L.) exposed to surfaces treated with methoxyfenozide

Electroantennogram (EAG) measurements were recorded from the antennae of male and female codling moth, Cydia pomonella L., to determine whether adult moths exposed to surfaces treated with the ecdysteroid agonist methoxyfenozide experience a decline in their antennal reception and thus olfactory sensitivity. Such a phenomenon would offer a possible mechanism for the previously reported decreased responsiveness from moths treated with methoxyfenozide to pheromone‐ and plant volatile‐based monitoring lures. Mean EAG data revealed that the antennae from methoxyfenozide‐treated male moths appear to be just as sensitive to various doses of synthetic codlemone as the antennae from the control and surfactant‐treated moths, but they appeared to be less sensitive to the pheromone component 12OH (collected from female effluvia) than the control male antennae. Mean male EAG responses to the pheromone components E8,E10‐12Al and codlemone collected from methoxyfenozide‐treated females were significantly less than the responses towards those two pheromone components collected from the control and surfactant‐treated females. Female moth exposure to methoxyfenozide did not negatively impact the sensitivity of female antennae to the plant volatile pear ester, but it did towards the apple volatile butyl hexanoate. Data from this study show that adult C. pomonella exposure to methoxyfenozide‐treated surfaces does appear to negatively impact, in a minor way, the (i) olfactory sensitivity (or detection) of male antennae towards some components of the female sex pheromone, (ii) the female antennal sensitivity towards a key apple volatile and (iii) the attractiveness of female pheromone effluvia.     

Investigation of a sex pheromone for the oriental cockroach,Blatta orientalis

A sex pheromone for adult male oriental cockroaches Blatta orientalis was isolated from the faeces of adult virgin female oriental cockroaches. It elicited a sexual response at 10 pg and 1 ng with B. orientalis and Periplaneta americana adult males, respectively. The site of production appers to be the crop, oesophagus, and proventriculus. Electroantennogram responses of male antennae toward the isolated pheromone were greater than those of the female antennae. The adult male oriental cockroach also responded to the American cockroach sex pheromone. The isolated pheromone with a mol. wt of 232 may be similar to one of the components of the American cockroach sex pheromone.     

Responses of Second-Instar Male Nymphs of Four Mealybug Species (Hemiptera: Pseudococcidae) to Conspecific and Heterospecific Female Sex Pheromones

The response of the late second-instar male nymphs of the mealybug species (Hemiptera, Pseudococcidae), Planococcus citri (Risso), Planococcus ficus (Signoret), Pseudococcus cryptus Hempel Nipaecoccus viridis (Newstead), to their conspecific and heterospecific female pheromone was studied. Males that exhibited the typical appearance of late-second-instar nymphs were tested. The male behavior was monitored soon after their exposure to the tested female sex pheromone in glass Petri dish arenas. Male nymph behavior toward the pheromone source was characterized based on their aggregation on the disks in the arena. Males of all four tested mealybug species were attracted to their conspecific female pheromone. By contrast, almost no interceptions of male nymphs with disks impregnated with a heterospecific female pheromone were observed. The mode of attraction of each of male nymphs of P. ficus, among most of the tested individuals (>80%), to the conspecific female sex pheromone, (S)-lavandulyl senecioate and or (S)-lavandulyl isovalerate, was the same as the mode of attraction latter on as adult. We suggest that by being attracted to the conspecific pheromone these males may direct themselves to a suitable pupation site near conspecific non-sibling mature females, thus preventing inbreeding. The repellency of heterospecific sex pheromone to males that are looking for a pupation site suggests that the latter try to avoid close contact with heterospecific females.     

Captures of oriental fruit moth, <Emphasis Type="Italic">Grapholita molesta</Emphasis> (Lepidoptera: Tortricidae), in traps baited with host-plant volatiles in Chile

Studies in Australia and China identified host-plant volatile blends from peach and pear that captured relatively high numbers of Grapholita molesta (Busck). To determine if these blends are attractants in other countries and relative to each other, the two host-plant blends, a laboratory blend identified in Switzerland, and a new “total blend” made by mixing components of all three blends, were field-tested in Chile for the first time. The same solvent type, concentrations, and dispensers as in the original studies, plus an additional concentration and solvent, were used. Only the Swiss blend at the low n-hexane concentration captured significantly more males than the solvent traps, albeit in very low numbers (1.46 ± 1.46, mean ± SEM males/trap/week). Furthermore, host-plant blends decreased male captures in sex pheromone traps, and the effect was dose-dependent for the Chinese and total blends. A laboratory flight tunnel test confirmed the lack of G. molesta male response to the Australian, Chinese, and Swiss plant blends. In the flight tunnel, however, the males responded sooner and in higher numbers to mixtures of sex pheromone with host-plant blends than they did to the sex pheromone alone.     

Changes in the amount of the diapause factor in the subesophageal ganglion during development of the silkworm, Bombyx mori

Changes in the diapause factor content of the subesophageal ganglion of the silkworm were examined from the viewpoint of sex and voltinism differences. The diapause factor content was comparatively low from the 4th larval ecdysis to the larval-pupal ecdysis, irrespective of sex and voltinism. However, remarkable changes occurred after the larval-pupal ecdysis. An interesting finding is that the diapause factor content in the female diapause egg-producer decreased gradually with pupal-adult development, but in the male diapause egg-producer as well as in the non-diapause egg-producer it increased up to the half-way stage in the pupal-adult development. This suggests that the diapause factor is not secreted from the subesophageal ganglion of the male pharate adult of the diapause egg-producer. This finding was confirmed by implantation experiments using brain-subesophageal ganglion complex or only subesophageal ganglion.     

Concurrent modulation of neuronal and behavioural olfactory responses to sex and host plant cues in a male moth

Mating has profound effects on animal physiology and behaviour, not only in females but also in males, which we show here for olfactory responses. In cotton leafworm moths, Spodoptera littoralis, odour-mediated attraction to sex pheromone and plant volatiles are modulated after mating, producing a behavioural response that matches the physiological condition of the male insect. Unmated males are attracted by upwind flight to sex pheromone released by calling females, as well as to volatiles of lilac flowers and green leaves of the host plant cotton, signalling adult food and mating sites, respectively. Mating temporarily abolishes male attraction to females and host plant odour, but does not diminish attraction to flowers. This behavioural modulation is correlated with a response modulation in the olfactory system, as shown by electro-physiological recordings from antennae and by functional imaging of the antennal lobe, using natural odours and synthetic compounds. An effect of mating on the olfactory responses to pheromone and cotton plant volatiles but not to lilac flowers indicates the presence of functionally independent neural circuits within the olfactory system. Our results indicate that these circuits interconnect and weigh perception of social and habitat odour signals to generate appropriate behavioural responses according to mating state.     

Low doses of a neonicotinoid insecticide modify pheromone response thresholds of central but not peripheral olfactory neurons in a pest insect

Insect pest management relies mainly on neurotoxic insecticides, including neonicotinoids, leaving residues in the environment. There is now evidence that low doses of insecticides can have positive effects on pest insects by enhancing various life traits. Because pest insects often rely on sex pheromones for reproduction, and olfactory synaptic transmission is cholinergic, neonicotinoid residues could modify chemical communication. We recently showed that treatments with different sublethal doses of clothianidin could either enhance or decrease behavioural sex pheromone responses in the male moth, Agrotis ipsilon. We investigated now effects of the behaviourally active clothianidin doses on the sensitivity of the peripheral and central olfactory system. We show with extracellular recordings that both tested clothianidin doses do not influence pheromone responses in olfactory receptor neurons. Similarly, in vivo optical imaging does not reveal any changes in glomerular response intensities to the sex pheromone after clothianidin treatments. The sensitivity of intracellularly recorded antennal lobe output neurons, however, is upregulated by a lethal dose 20 times and downregulated by a dose 10 times lower than the lethal dose 0. This correlates with the changes of behavioural responses after clothianidin treatment and suggests the antennal lobe as neural substrate involved in clothianidin-induced behavioural changes.     

Circadian mating activity and effect of pheromone pre-exposure on pheromone response rhythms in the moth Spodoptera littoralis

Mating in moths is generally mediated by female-produced sex pheromones. Mating activity, female pheromone production/release and male pheromone responsiveness all show diurnal variations in many species. We found that the response of the male Egyptian cotton leafworm, Spodoptera littoralis, to sex pheromone gland extracts showed a diel rhythm in olfactometer tests, and the variation was persistent for at least 1 day in constant darkness. High male response to sex pheromone was correlated in time with high mating and locomotor activity. Male S. littoralis, maintained in constant darkness and exposed to pheromone gland extracts on a daily basis, showed an induced temporal variation in response after several days, in contrast to unexposed males. This suggests that in the absence of other external zeitgebers, exposure to sex pheromone may function to synchronise circadian behavioural rhythms in male moths. The daily rhythm in mating activity in S. littoralis is also shown to be persistent for at least 2 days in constant darkness. Pairs mated significantly less when either the male or female had been raised in a light:dark cycle 10 h out of phase, indicating that the proposed circadian rhythm in mating activity is composed of rhythmic mating preference/ability in both sexes.     

温度对二化螟性信息素通讯的影响

环境因素影响昆虫两性间的化学通讯,也影响性信息素技术的田间防治效果.本文探讨了温度对二化螟雌蛾性信息素产生以及雄蛾对性信息素触角电位反应的影响,以期为田间二化螟的性信息素防治提供指导.在二化螟蛹期和成虫期进行不同温度处理(15、20、25、30和35 ℃),然后利用气相色谱仪(GC)分析雌蛾性腺内各性信息素组分的含量及比例,同时利用触角电位仪(EAG)测定雄蛾对性信息素组分的电生理反应.结果表明: 25 ℃处理中雌蛾性腺内3个性信息素组分(Z9-16:Ald、Z11-16:Ald和Z13-18:Ald)的含量均显著高于其他温度处理(15、20和30 ℃),且25 ℃处理中Z13-18:Ald的相对比例也显著低于其他温度处理.就雄蛾对性信息素的敏感性而言,对3种性信息素单一组分及特定比例混合物的EAG反应在15～25 ℃间没有显著差异,但在25～35 ℃间(Z13-18:Ald在30～35 ℃间)随温度升高呈下降趋势,且30 ℃较25 ℃显著降低,35 ℃较30 ℃又显著降低.综合分析认为,二化螟性信息素通讯的适宜温度为20～25 ℃,温度过高或过低均不利于二化螟两性间的正常化学通讯.研究结果为二化螟性信息素防治技术的合理应用及极端温度条件下害虫种群发生的预测预报,提供了重要参考.     

Perception of volatiles associated with sex and food by different adult forms of the black bean aphid, Aphis fabae

Abstract. Electroantennograms (EAGs) were recorded from adult male and asexual forms (winged and wingless virginoparae and gynoparae) of the black bean aphid, Aphis fabae , during stimulation with two sex pheromone components, (+)-(4a S ,7 S ,7a R )-nepetalactone and (-)-(1 R ,4a S ,7 S ,7a R )-nepetalactol, as well as six plant volatiles, i.e. ( E )-2-hexenal, ( E )-2-hexenol-1, ( Z )-3-hexenol-1, ( Z )-3-hexenyl acetate, hexanal and allyl isothiocyanate. The male antennae are 1000-10,000 times more sensitive to nepetalactol and nepetalactone than to the plant compound ( E )-2-hexenal. Besides this marked difference of EAG peak responses in males, the EAG rise and decay are slower for the sex pheromone components. Males are also much more sensitive to the sex pheromone components than asexual females. This high sensitivity correlates with a predominance of antennal secondary rhinaria, the major sites of pheromone perception in the male. However, it is the primary rhinaria on the antennae of the wingless asexual females that are responsible for pheromone perception. Male antennae are as responsive as the asexual female antennae to the plant volatiles. The specialization of the male for mate location is discussed.     

Environmental conditions terminate reproductive diapause and influence pheromone perception in the long‐lived moth Caloptilia fraxinella

Reproductive diapause enables long‐lived insects to time mating with environmental conditions suitable for offspring development. Plasticity in the perception of pheromones used in sexual communication may enable mate‐finding at the appropriate time of year. The moth Caloptilia fraxinella (Ely) (Lepidoptera: Gracillariidae) undergoes a 9‐month reproductive diapause, during which the male response to pheromone is plastic and is highest during the period of reproductive activity. The mechanisms controlling this pheromone response plasticity are not well‐understood, and the aim of the present study is to determine the main factors involved. In the present study, the impact of temperature, photoperiod, juvenile hormone analogue (JHA) and adult nutrition on diapause termination are tested using electroantennogram (EAG) and behavioural response to pheromone in male C. fraxinella. Eclosion in a state of reproductive diapause occurs in most males; diapause is maintained under short‐day or cool conditions indoors, or under natural conditions outdoors. Exposure to long‐day, warm conditions over a period of 4 weeks causes a small number of males to become behaviourally responsive to pheromone; a larger number of males become behaviourally responsive over a period of 3 months of post‐eclosion. Treatment with a JHA impacts male EAG and the behavioural response to pheromone during the period of reproductive diapause. A carbohydrate food source is not required by reproductively active adult male C. fraxinella to respond to pheromone and express mate location behaviours. The main factors involved in controlling male pheromone response plasticity and the implications of these factors for the C. fraxinella population in its expanded range are discussed.     

Anatomical site of pheromone accumulation and temporal pattern of pheromone emission in the ambrosia beetle Megaplatypus mutatus

Megaplatypus mutatus (Chapuis) is a native South American ambrosia beetle that attacks live hardwood trees (e.g. Populus spp.), causing important economic losses to commercial plantations. Male beetles release the main components of the sex pheromone, namely (+)‐6‐methyl‐5‐hepten‐2‐ol [(+)‐sulcatol, or retusol] and 6‐methyl‐5‐hepten‐2‐one (sulcatone), when colonizing suitable hosts. The hindgut is shown to be the anatomical site of pheromone accumulation within males, the enantiomeric composition of sulcatol in this tissue is 99%‐(+) and sulcatol is first detectable in this tissue on days 1–2 after gallery initiation. Peak accumulation of sulcatol occurs on days 5–12 after gallery initiation. Trace quantities of sulcatone are also observed during the same period. Both pheromone components are present in male emissions from three host species (Populus×canadensis, Populus alba and Casuarina stricta) between days 2 and 12 after gallery initiation, although sulcatone is always present in low concentrations. The temporal patterns of sulcatol and sulcatone accumulation or storage in male M. mutatus correspond to the temporal patterns of emission.     

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.     

Production and evolution pattern of “fruity smell” aggregation pheromones in genus Drosophila

Insects produce pheromones to serve a range of ecological functions throughout their lifetime. The chemical composition, production pattern, and interspecies specificity provide information for carrying out their function and biological significance. Several species of Drosophila produce a class of volatile esters considered as “fruity smells”; however, the production pattern and ecological functions of these “fruity smell” volatiles in genus Drosophila are poorly understood. Here, using the headspace solid-phase microextraction (SPME) method, we tested the production pattern of volatile pheromones in Drosophila immigrans and factors that possibly affected pheromone production, including mating, feeding conditions, age of adult flies, and geographical distribution. We also explored the evolution and production pattern of volatile pheromones in 14 species of genus Drosophila. Our result showed that male D. immigrans adult flies produce three male-specific volatile ester pheromones, which are also considered as “fruity smell” chemicals, in a relatively stable pattern. In addition, a series of “fruity smell” ester pheromones with similar structure and chemical properties were found to appear in the species of D. virilis and D. immigrans species group, but not in the species of D. melanogaster species group. The ester volatile pheromone production of male flies has a correspondence with the female's demand for host plants. Integrating the production and evolution pattern of these volatile chemicals, we inferred the interaction between insects and host plants reflected in the Drosophila “fruity smell” pheromones.     

Location,morphology and histology of sex pheromone glands of the female gypsy moth, Lymantria dispar (L.)

Scanning electron microscopy, histology and a male wing fanning bioassay were used in this study to locate the sex pheromone-producing glands of the female gypsy moth, Lymantria dispar. When exposed to female sex pheromone, adult males exhibit a strong wing fanning behaviour prior to take off. We found that adult males showed positive response to calling females and to tissue extract from both dorsal and ventral portions of the intersegmental membrane between 8th and 9th-abdominal segments. A typical male response usually starts with elevation of antennae, movement of head in different directions, walking, wing fanning and onset of search flight. Histological and scanning electron microscopic studies suggested that the sex pheromone glands are located on the dorsal and ventral aspects of the intersegmental membrane. The glands appear as two highly convoluted integumentary areas with hypertrophied glandular epidermal cells.     

Sound production in Scolytidae: Female sonic stimulus of male pheromone release in two Dendroctonus beetles

The hypothesis that female sonic stimulus may evoke male pheromone release in a behavioural interaction analogous to the known male sonic stimulus of female pheromone release, was confirmed in Dendroctonus pseudotsugae, and also in D. brevicomis. In both species known male-produced substances collected from males stimulated by recorded female stridulation were identified by coupled gas chromatography/mass spectrometry. In a second test with D. pseudotsugae, male pheromone release during recorded female stridulation was evident in the change of male stridulation from the simple attractant chirp to the interrupted chirp, which is known to result from a medium concentration of 3,2-MCH. Also, the D. pseudotsugae male attractant chirp was synthesised with an electronic pulse generator and used to evoke pheromone release. It is concluded that the antiaggregative pheromone of this species is released by each sex at the sonic stimulus of the other sex.