Detection of short pure-tone stimuli in the noctuid ear: what are temporal integration and integration time all about?

The interception of a pheromone filament induces flying moths to surge briefly nearly straight upwind; in the absence of pheromone moths cease upwind progress and zigzag crosswind. We tested males of the almond moth, Cadra cautella (Lepidoptera, Pyralidae), in a low-turbulence wind tunnel in wind velocities of 20, 40 and 80 cm s⁻¹. A mechanical pulse generator was set to produce plumes either with same pheromone pulse frequency (pulse generation frequency of 2.9 Hz, interpulse distances from 7 cm to 28 cm) or plumes with same interpulse distance across the three wind velocities (interpulse distance of 14 ± 2 cm, pulse generation frequency of 1.7–5.0 Hz). In plumes of similar pulse frequency, the faster the speed of the wind the slower the ground speed of flight. However, in plumes of similar interpulse distance, ground speed remained relatively constant independent of the wind speed. A `realized' frequency of pulse interception for males flying along the various combinations of pulse frequencies and wind velocities was calculated using the males' average airspeed and the spatial distribution of pheromone pulses in the plume. Realized frequency of pulse interception ranged from 1.3- to 3.0-fold higher than the frequency of pulse generation. The flight tracks of males reflected the regime of realized pulse interception. These results suggest that upwind flight orientation of male C. cautella to pheromone in different wind velocities is determined by the flux of filament encounter. Accepted: 3 September 1997     

The Effect of Abiotic Factors on the Male Mate Searching Behavior and the Mating Success of Aphidius ervi (Hymenoptera: Aphidiidae)

The effect of wind speed and distance from the source on the male response of the aphid parasitoid, Aphidius ervi (Hymenoptera: Aphidiidae), to a pheromone source was studied in a wind tunnel. The number of males taking flight, entering the plume and successfully reaching the source, decreased at wind speeds >50 cm/s. Furthermore, the proportion of those attempting upwind flight that fell to the ground increased with increasing wind speed. In contrast, distance from the source had no significant effect on any of the parameters examined. While male flight behavior was significantly reduced at 70 cm/s, some males walked to the source when there was a bridge connecting the pheromone source and the release platform. This suggests that ambulatory behavior could be a significant component of male mate searching in A. ervi when wind conditions are too strong for upwind flight. The possible effects of variation in atmospheric pressure on male flight behavior to the long distance pheromone, as well as to the short distance one, were also investigated. No significant effects of atmospheric pressure were observed. These findings differ significantly from those previously reported for another aphid parasitoid, A. nigripes, and the reasons for such differences are discussed.     

Effects of Wind Speed and Atmospheric Pressure on Mate Searching Behavior in the Aphid Parasitoid Aphidius nigripes (Hymenoptera: Aphidiidae)

The effects of wind speed and atmospheric pressure on male mate searching behavior, modulated by a female sex pheromone, were investigated in the aphid parasitoid Aphidius nigripes (Hymenoptera: Aphidiidae). Male A. nigripe generally did not reach females at wind speeds of 100 cm/sec, as the majority of individuals taking flight in the pheromone plume (81.8%) were unable to sustain upwind flight. At lower wind velocities, male responsiveness to females generally decreased with distance from the source. However, wind speeds approaching the upper threshold (100 cm/sec) tended to eliminate this distance effect. Therefore, there appears to be a trade-off between the need for higher wind speeds to detect the pheromone source from long distances, and a reduction in male flight capacity as wind velocity increases. Our results also indicate that chemical communication in A. nigripes could be affected by variations in atmospheric pressure, as we observed a relationship between pressure fluctuations in the 24 hr prior to testing and male responsiveness to females. The importance of these abiotic factors on mate searching behavior is discussed within the context of the reproductive biology of A. nigripes.     

二齿新蚤和方形黄鼠蚤松江亚种侵袭与离开宿主习性的实验研究

本文对二齿新蚤和方形黄鼠蚤松江亚种侵袭与离开宿主的习性进行了实验研究,结果：(1)蚤攻击宿主距离平均在2cm以内,最大攻击距离不超过10cm。在有效侵袭范围内,蚤与宿主距离近时较距离远时吸血蚤数增多。(2)蚤对宿主的侵袭程度雌蚤大于雄蚤,繁殖蚤大于新羽化蚤,二齿新蚤大于方形黄鼠蚤松江亚种,对小白鼠大于对达乌尔黄鼠。(3)宿主死后一定时间内,仍有部份蚤侵袭其尸体。 死亡时间越长侵袭蚤数越少,呈logY=a-blogX型曲线。(4)蚤离开宿主时间呈偏态分布,温度越高偏态分布越明显。蚤离开死鼠和离开活鼠所需时间很接近。 二齿新蚤和方形黄鼠蚤松江亚种离开宿主时间几乎相同。二种蚤离开宿主平均时间与环境温度呈负相关,且呈曲线关系。     

Moth sex pheromone adsorption to leaf surface: bridge in time for chemical spies

Abstract When brussels sprouts leaves were exposed to air passed over a single calling Mamestra brassicae moth, sex pheromone was adsorbed onto the leaf surface to such an extent that it subsequently elicited behavioural responses in conspecific male moths, as well as in female Trichogramma evanescens , egg parasitoids of M. brassicae. Male moths responded to odour-treated leaves over a short distance (c. 5 cm) in a wind tunnel, but were not attracted at a distance of 1 m. In contrast, sex pheromone adsorbed to and re-released from the glass wall of a wind tunnel compartment attracted male moths from 1 m. Trichogramma's locomotory behaviour on leaves is described. On leaves previously exposed to air passed over a calling female moth, Trichogramma wasps stayed significantly longer on leaves treated with clean air or air passed over a non-calling female moth. In addition, wasps spent relatively more time along the margin and on the leaf underside on treated leaves compared with control leaves. All effects persisted for at least 4 h, and after 24 h treated leaves still increased wasp residence times. These results are the first example of responses of male moths to adsorbed airborne sex pheromone originating from a single female moth. They further suggest how sex pheromone released by nocturnal moths may function as a kairomone for diurnally foraging parasitoids.     

Host plant volatiles serve to increase the response of male European grape berry moths, <Emphasis Type="Italic">Eupoecilia ambiguella,</Emphasis> to their sex pheromone

The European grape berry moth is an important pest in vineyards. Males respond to the female-produced sex pheromone released from a piezo nebulizer in a dose-dependent manner in a wind tunnel: <50% arrive at the source at 5–50 pg/min (underdosed), 80% arrive at 100 pg/min to 10 ng/min (optimal) and <20% arrive at 100 ng/min (overdosed). Males responding to overdosed pheromone show in flight arrestment at 80 cm from the source. Host plant chemostimuli for Eupoecilia ambiguella increase the responses of males to underdosed and overdosed pheromone. (Z)-3-hexen-1-ol, (+)-terpinen-4-ol, (E)-β-caryophyllene and methyl salicylate released with the underdosed pheromone cause a significant increase in male E. ambiguella flying to the source. Time–event analysis indicates a positive correlation between faster activation and probability of source contact by the responding males. The four host plant compounds added to the overdosed pheromone permitted males to take off faster and with a higher probability of flying to the source. This suggests that perception of host plant products with the sex pheromone facilitates male E. ambiguella to locate females on host plants, lending credence to the hypothesis that plant products can signal rendezvous sites suitable for mating. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of velocity-specific training on rate of velocity development, peak torque, and performance

Little is known about the velocity-specific adaptations to training utilizing movement velocities in excess of 300 degrees x s(-1). The purpose of this investigation was to determine the effects of 4 weeks of slow (60 degrees x s(-1)) vs. fast (400 degrees x s(-1)) velocity training on rate of velocity development (RVD), peak torque (PT), and performance. Twenty male kinesiology students (22.0 years +/- 2.72; 178.6 cm +/- 7.1; 82.7 kg +/- 15.5) were tested, before and after 4 weeks of training, for PT production, RVD (at 60, 180, 300, 400, and 450 degrees x s(-1)), standing long jump (SLJ) distance, and 15- and 40-m sprint times. All participants underwent 8 training sessions, performing 5 sets of 5 repetitions of simultaneous, bilateral, concentric knee extension exercises on a Biodex System 3 isokinetic dynamometer at either 60 degrees or 400 degrees per second. Two 5 (speed) x 2 (time) x 2 (group) multivariate repeated measures analyses of variance revealed no significant differences between groups on any measure. Therefore, the groups were collapsed for analysis. There was a significant (p < 0.05) main effect for RVD by time and SLJ distance by time (pre- 227.1 cm +/- 21.2; post- 232.9 cm +/- 20.7) but no significant change in PT or 15- or 40-m sprint times. These results offer support for the suggestion that there is a significant neural adaptation to short-term isokinetic training performed by recreationally trained males, producing changes in limb acceleration and performance with little or no change in strength. Because results were independent of training velocity, it appears as though the intention to move quickly is sufficient stimulus to achieve improvements in limb RVD. Changes in SLJ distance suggest that open kinetic chain training may benefit the performance of a closed kinetic chain activity when movement pattern specificity is optimized.     

Real-time measurement of pheromone release from individual female moths and synthetic dispensers in a wind tunnel by recording of single receptor-neurone responses

Measurements of single neurone activity in the peripheral pheromone receptors of male Agrotis segetum (Denis & Schiffermüller) (Lepidoptera: Noctuidae) were performed in a wind tunnel using a portable electrophysiological recording unit. Filter paper and rubber septa loaded with synthetic sex pheromone, as well as individual conspecific female glands, were used as pheromone sources. Recordings, up to 3 h long, were analysed for temporal variation in spiking activity. The recordings were performed 2 m downwind of the source, where the pheromone plume had a width of approximately 12 cm, as could be measured with the single cell preparations. The system allowed reliable measurements of relative pheromone concentration with a 20-s time resolution. The release rate from rubber septa loaded with pheromone was more or less constant over time, whereas the release rate from filter paper loaded with pheromone decreased to one tenth of the initial value within 6 min from the application of the pheromone. The release of pheromone from female pheromone glands was pulsed with an interval of 2–10 min between bursts. This pulsing was not caused by retraction of the gland, as the glands were forcibly extruded during the entire experiment, but should reflect variation in transport of pheromone to the gland surface and subsequent release. The demonstrated stability of the preparations using tungsten electrodes, the reliable monitoring of female-produced pheromone plumes at several metres distance, and the time resolution obtained are important steps towards field monitoring of natural pheromone plumes, as well as pheromone concentration and distribution in applications for mating disruption.     

Host instar, body size and fitness in the koinobiotic parasitoid Aphidius nigripes

In aphidiine parasitoids, resources for growth and adult body size increase with host instar used by ovipositing females, but the fitness consequences of body size on fitness are poorly documented. We compared the fitness of male and female A. nigripesadults that varied in size as a consequence of developing in different instars of their host Macrosiphum euphorbiae. When reproductive fitness was measured without considering time, female wasps from small and large hosts performed similarly, contributing 125–175 foundresses plus 100–180 sons to the next generation. However, when expressed as the innate capacity for increase (r _m), female fitness correlated with host-induced variation of wasp size, indicating that micropopulations initiated by large wasps would increase faster. In a wind-tunnel, a sex pheromone plume from large female wasps induced more males to fly upwind when released at a distance of 50 cm downwind than small females, indicating that large females were sexually more attractive. With respect to male body size effects on fitness, large individuals performed similar to small ones, whether fitness was measured by lifetime mating frequency, fertile inseminations, or proportion of daughters among progeny born from their mates. When young naive males of unequal size were directly competing for mating with a virgin female, small and large males had equal mating success, and large individuals were no more successful than small ones at displacing a competitor already positioned on a receptive female. In a wind-tunnel test where males were scored on their ability to reach a female pheromone source, small and large males were equally affected by wind speed but reached the source located 50 cm downwind in equal proportions, suggesting similar capacity for finding mates by flying upwind. Our results indicate that despite host resources not being fixed at the time of attack for the koinobiont A. nigripes, fitness consequences of resource limitation by the mother may be perceived to be greater for daughters than sons, which would explain male-biased sex ratio in early-instar hosts.     

Grasshopper sexual pheromone: a component of the defensive secretion in Taeniopoda eques

ABSTRACT. Both male and female Taeniopoda eques (Burmeister) emit a defensive secretion from their thoracic spiracles. The secretion from mature females was found to act as a sex pheromone, eliciting mating behaviour in males. Females became chemically attractive to males about 16–18 days after eclosion. The antennae are shown to be the site of pheromone reception in males. Feeding upon natural host plants was not a prerequisite for pheromone production.     

植物气味化合物与斜纹夜蛾性信息素的协同作用

为提高现有性信息素对雄蛾的引诱活性, 本研究通过大量的田间试验探索植物气味化合物与斜纹夜蛾Spodoptera litura性信息素（顺9, 反11-十四碳二烯乙酸酯∶顺9, 反12-十四碳二烯乙酸酯=10∶1）的协同作用机制。从斜纹夜蛾寄主植物和花的气味化合物中, 选择9种有代表性的化合物, 并以一定剂量分别加入到斜纹夜蛾性信息素诱芯中, 在田间测试对雄蛾的引诱活性。结果表明: 在测试的9种植源性化合物中, 发现一定剂量(每个诱芯加入0.4 mg)的苯乙醛(PAA), 具有显著提高斜纹夜蛾性信息素的引诱作用, 而高剂量的苯乙醛则强烈抑制性信息素的引诱活性; 此外, 其他各种浓度的测试化合物或混合物对性信息素则没有统计上显著的增效作用。不同剂量的苯乙醛单个化合物及各种植物气味化合物组成的混合物对斜纹夜蛾也有微弱的引诱作用。苯乙醛必须要与性信息素的完整组分(以10∶1比例混合的顺9, 反11-十四碳二烯乙酸酯和顺9反, 12-十四碳二烯乙酸酯)混合才能起作用, 缺少顺9, 反12-十四碳二烯乙酸酯则没有引诱活性。本研究证明, 苯乙醛作为理想的性信息素诱芯增效剂, 可应用于建立更理想的斜纹夜蛾性信息素诱杀技术, 对性诱害虫防治和测报具有应用价值。     

Spatial discrimination of pheromones and behavioural antagonists by the tortricid moths Cydia pomonella and Adoxophyes orana

Male moths responding to their species-specific sex pheromone, may cease their upwind flight when pheromone components of sympatric species are added to the mixture. The interspecific interaction between the pheromone response of the tortricid moths Cydia pomonella and Adoxophyes orana was investigated in field-trapping and wind-tunnel studies. Addition of the A. orana pheromone [(Z9)-tetradecenylacetate and (Z11)-tetradecenylacetate] to a source containing the C. pomonella pheromone [(E8, E10)-dodecadienol] resulted in a significant inhibition of attraction by male C. pomonella. It is demonstrated that this behavioural antagonist for C. pomonella must be emitted from the same point source to induce this inhibitory effect. A spatial separation of the two interspecific pheromones (at 14 cm, 5 cm and 0.5 cm crosswind) restored the attraction of the conspecific pheromone for male C. pomonella. In contrast to C. pomonella, male A. orana were not inhibited by point sources releasing both the C. pomonella and A. orana pheromone. We suggest that the discrepancy in the interspecific pheromone interaction between these two tortricids can be explained if we consider the evolutionary ecology of interspecific pheromone communication in C. pomonella and A. orana. Accepted: 18 July 1999     

Metacestode-induced depression of the production of, and response to, sex pheromone in the intermediate host Tenebrio molitor

Hymenolepis diminuta infection of Tenebrio molitor is associated with an impairment of vitellogenesis and a reduction in host fecundity. In this communication the effect of infection upon an additional aspect of host reproduction, the initiation of mating behavior, has been examined. Copulatory release pheromone, extracted from control virgin females 6-7 days old, was shown to stimulate a positive mating response in 88% of 5- to 6-day-old control males; however, only a 56% response was elicited by pheromone from infected females. In addition, parasitization adversely effected male response to pheromone from control females. A significant (P less than 0.001) depression of copulatory response occurred in infected 6- to 7-day-old males (age of peak response) although this effect was not sustained in older beetles. The possibility that an endocrine interaction between metacestodes and host may mediate these effects is discussed in the light of our knowledge of the role of host juvenile hormone in controlling both pheromone production and vitellogenesis in T. molitor.     

Factors influencing adult male Grapholita molesta dispersal in commercial Malus and Prunus host crops

The oriental fruit moth (OFM), Grapholita molesta (Busck) (Lepidoptera: Tortricidae), has been an economic pest of apples (Malus spp.) and peaches (Prunus spp.; both Rosaceae) in the eastern USA since the 1930s. Successful management of G. molesta with insecticides requires careful timing of these products based on sex‐pheromone trap captures of male moths. In Pennsylvania, apple and peach orchards are often planted adjacently. Factors such as trap distance from release points, host crop, and wind were considered in relation to male moth movement within and between hosts. Our practical objective was to understand male G. molesta dispersal within and between these two hosts to achieve better management. Recapture of moths decreased over distance; the majority of moths were recaptured 50–100 m from the release point regardless of crop type. Male G. molesta released into commercial apple and peach orchards were most likely to remain in the crop of release and to not exhibit host shifts over the season, even after peach fruit were harvested. Our analyses of wind direction data with respect to recapture showed that the likelihood of recapture was similar regardless of the orientation of the traps relative to the mean wind direction at peak flight times. The apple and peach host crops appeared to affect male G. molesta dispersal to the extent that wind in orchards affects their perception of pheromone. Crop effects were most likely due to the traps closest to the release points being in the same crop as the release points.     

Le grégarisme chez Blaberus craniifer: Isolement et identification de la phéromone

In the gregarious cockroach, Blaberus craniifer, an aggregation pheromone is produced by all individuals (larvae and adults) except at ecdysis. The pheromone, secreted by the mandibular glands, was analysed by thin-layer and gas chromatography and by mass spectrometry. The mandibular glands secrete three major volatile products: undecane, tetradecane, and ethyl-caproate. The last component is of unknown significance, but a mixture of undecane and tetradecane ($\text{1}\text{1}$) reproduces all the effects of the natural pheromone. The distance of perception is short: 40 cm for the adults and old larvae and 10 cm for the first instar larvae. The threshold is about 0·2 ng for the adults and 0·4 ng for the first instar larvae. The mixture undecane and tetradecane is specific: it is not attractive for other gregarious cockroaches (many of which are devoid of mandibular glands), and other straight-chain saturated hydrocarbons are not attractive for B. craniifer.     

Host plant volatiles synergize responses of sex pheromone-specific olfactory receptor neurons in male <Emphasis Type="Italic">Helicoverpa zea</Emphasis>

Single-cell electrophysiological recordings were obtained from olfactory receptor neurons in antennal trichoid sensilla of male corn earworm, Helicoverpa zea. Spontaneous activity of the neuron specific for the major component ( Z)-11-hexadecenal, the conspecific female-emitted sex pheromone, was not affected by exposure to host plant volatiles. However, stimulations with binary mixtures of a threshold dosage of the pheromone component and increasing dosages of either linalool or ( Z)-3-hexenol significantly synergized the pheromone-specific neuron's firing rates compared with responses to the major pheromone component alone. Cross-adaptation studies confirmed that the enhanced impulses originated from the pheromone-component-tuned neuron. Because plant volatiles do not stimulate the pheromone-specific neuron when presented alone, the pheromone plus host odor blend would be interpreted as containing more pheromone than it actually does when processed by the pheromone-processing portion of the antennal lobe.     

Effects of different sex pheromone compositions and host plants on the mating behavior of two Grapholita species

The two congener species Grapholita molesta and Grapholita dimorpha share two major sex pheromone components: cis-8-dodecenyl acetate (Z8-12Ac) and trans-8-dodecenyl acetate (E8-12Ac). In fact, commercial sex pheromone lures composed of only these two major components attract the males of both species. In this study, we aimed to determine the reproductive isolation components of these two species by analyzing the effects of the minor sex pheromone components and host plants. First, different ratios of the two major sex pheromone components were greatly favored by either male species. Sex pheromone gland extracts of G. dimorpha contained a lesser proportion of Z8-12Ac than that of G. molesta. In the three (apple, pear, and peach) orchards investigated in this study, a larger number of G. molesta males were attracted to the 95:5 pheromone mixture (Z8-12Ac and E8-12Ac, respectively), while a larger number of G. dimorpha males were attracted to the 85:15 mixture. Second, there was a significant variation in male attractions in different host plants. G. molesta males were more attracted to the sex pheromone lure in the apple orchards than that in the pear and peach orchards. In contrast, G. dimorpha males were more attracted to the lures in the pear and peach orchards than that in the apple orchard. Third, the minor sex pheromone components were important for reproductive isolation. Among the four minor components tested, addition of (Z)-8-dodecenol (Z8-12OH) to the major sex pheromone components significantly suppressed male attraction in G. dimorpha and slightly elevated male attraction in G. molesta. The discriminating effect of Z8-12OH was further validated using male electroantennogram analysis. These results suggest that reproductive isolation between two congeners can be achieved by variations in the minor sex pheromone components and in the host plants, as well as by changes in the ratio of the two major components.     

Sex pheromone and visual trap interactions in mate location strategies and aggregation by host-alternating aphids in the field

Abstract. Field observations were made on the responses of males and gynoparae of three host-alternating aphid species, the blackberry-cereal aphid, Sitobion fragariae (Walker), the bird cherry-oat aphid, Rhopalosiphum padi (L.) and the damson-hop aphid, Phorodon humuli (Schrank) to species-specific sex pheromones released from transparent and coloured water traps.Pheromone traps caught significantly more males than did control traps without pheromone, whereas transparent, light green, yellow and orange traps caught most insects.Measurements of the distance over which sex pheromones function indicated that male P.humuli detect the pheromone 2–6 m from the source and can fly upwind to a source in wind speeds of 0.7 m s^-1.In all three species significantly more gynoparae were caught in pheromone traps than in control traps, suggesting that pheromone released by adult sexual females may assist late-flying gynoparae to locate a suitable host plant on which to deposit their progeny.The response is relatively stronger for males than gynoparae, but the pheromones appear to act as both sex and aggregation pheromones.