Intraspecific acoustic communication and mechanical sensitivity of the tympanal ear of the gypsy moth Lymantria dispar

Tympanal ears of female gypsy moths Lymantria dispar dispar (L.) (Lepidoptera: Erebidae: Lymantriinae) are reportedly more sensitive than ears of conspecific males to sounds below 20 kHz. The hypothesis is tested that this differential sensitivity is a result of sex‐specific functional roles of sound during sexual communication, with males sending and females receiving acoustic signals. Analyses of sounds produced by flying males reveal a 33‐Hz wing beat frequency and 14‐kHz associated clicks, which remain unchanged in the presence of female sex pheromone. Females exposed to playback sounds of flying conspecific males respond with wing raising, fluttering and walking, generating distinctive visual signals that may be utilized by mate‐seeking males at close range. By contrast, females exposed to playback sounds of flying heterospecific males (Lymantria fumida Butler) do not exhibit the above behavioural responses. Laser Doppler vibrometry reveals that female tympana are particularly sensitive to frequencies in the range produced by flying conspecific males, including the 33‐Hz wing beat frequency, as well as the 7‐kHz fundamental frequency and 14‐kHz dominant frequency of associated clicks. These results support the hypothesis that the female L. dispar ear is tuned to sounds of flying conspecific males. Based on previous findings and the data of the present study, sexual communication in L. dispar appears to proceed as: (i) females emitting sex pheromone that attracts males; (ii) males flying toward calling females; and (iii) sound signals from flying males at close range inducing movement in females, which, in turn, provides visual signals that could orient males toward females.     

Acoustic communication in Okanagana rimosa (Say) (Homoptera: Cicadidae)

The cicada Okanagana rimosa (Say) has an acoustic communication system with three types of loud timbal sounds: (i) A calling song lasting several seconds to about 1 min which consists of a sequence of chirps at a repetition rate of 83 chirps per second. Each chirp of about 6 ms duration contains 4-5 pulses. The sound level of the calling song is 87-90 dB SPL at a distance of 15 cm. (ii) An amplitude modulated courtship song with increasing amplitude and repetition rate of chirps and pulses. (iii) A protest squawk with irregular chirp and pulse structure. The spectra of all three types are similar and show main energy peaks at 8-10 kHz. Only males sing, and calling song production is influenced by the songs of other males, resulting in an almost continuous sound in dense populations. In such populations, the calling songs overlap and the temporal structure of individual songs is obscured within the habitat. The calling song of the broadly sympatric, closely related species O. canadensis (Provander) is similar in frequency content, but distinct in the temporal pattern (24 chirps per second, 24 ms chirp duration, eight pulses per chirp) which is likely important for species separation in sympatric populations. The hearing threshold of the auditory nerve is similar for females and males of O. rimosa and most sensitive at 4-5 kHz. Experiments in the field show that female phonotaxis of O. rimosa depends on parameters of the calling song. Most females are attracted to calling song models with a 9 kHz carrier frequency (peak frequency of the calling song), but not to models with a 5 kHz carrier frequency (minimum hearing threshold). Phonotaxis depends on temporal parameters of the conspecific song, especially chirp repetition rate. Calling song production is influenced by environmental factors, and likelihood to sing increases with temperature and brightness of the sky. Correspondingly, females perform phonotaxis most often during sunny conditions with temperatures above 22 degrees C. Non-mated and mated females are attracted by the acoustic signals, and the percentage of mated females performing phonotaxis increases during the season.     

摄食声对草鱼幼鱼的诱集作用

以草鱼(Ctenopharyngodon idellus)幼鱼为实验对象, 进行了声音播放实验, 旨在探究草鱼幼鱼对水下录制的草鱼摄食浮萍声音(简称摄食声)的行为反应。以不播放声音的草鱼鱼群作为对照, 探讨了4 种单频音(500、1000、2000和3000 Hz)和摄食声对草鱼游泳行为和在水槽内的分布的影响。结果表明: 在播放单频音时, 3min内草鱼的趋音游泳速度和逗留时间与对照组无显著性差异(P>0.05); 在播放摄食声时, 3min内草鱼的趋音游泳速度和逗留时间显著高于4种单频音组和对照组(P<0.05); 在播放单频音时, 20min内草鱼的平均游泳速度、水槽内的分布和趋音率与对照组无显著性差异(P>0.05); 在播放摄食声时, 20min内草鱼的平均游泳速度和趋音率都显著高于4种单频音组和对照组(P<0.05)。草鱼摄食声对草鱼幼鱼有诱集作用, 为声音诱鱼技术研究提供了科学依据。     

Nanometre-range acoustic sensitivity in male and female mosquitoes

Johnston's sensory organ at the base of the antenna serves as a movement sound detector in male mosquitoes, sensing antennal vibrations induced by the flight sounds of conspecific females. Simultaneous examination of acoustically elicited antennal vibrations and neural responses in the mosquito species Toxorhynchites brevipalpis has now demonstrated the exquisite acoustic and mechanical sensitivity of Johnston's organ in males and, surprisingly, also in females. The female Johnston's organ is less sensitive than that of males. Yet it responds to antennal deflections of +/- 0.0005 degrees induced by +/- 11 nm air particle displacements in the sound field, thereby surpassing the other insect movement sound detectors in sensitivity. These findings strongly suggest that the reception of sounds plays a crucial role in the sensory ecology of both mosquito sexes.     

Broadcasts of wing-fanning vibrations recorded from calling male Ceratitis capitata (Diptera: Tephritidae) increase captures of females in traps

Female Mediterranean fruit flies, Ceratitis capitata (Wiedemann), from the sterile-male rearing facility in El Pino, Guatemala, were exposed to broadcasts of wing-fanning vibrations recorded from males engaged in calling behavior to investigate the feasibility of developing a female-selective acoustic trap. The recorded signals had frequent amplitude fluctuations and peak frequencies approximately 350 Hz, typical of signals observed in previous studies of Mediterranean fruit fly acoustic behavior. Females did not exhibit long-distance phonotaxis, but remained near a speaker significantly longer when the sounds were broadcast at 103-107 dB than when the speaker was silent. In addition, significantly higher percentages of females were captured by yellow adhesive traps next to a broadcasting speaker than by traps next to a silent mimic. Additional bioassays were conducted with synthetic, 350-Hz tones produced by a thermoacoustic tube as well as with silent mimics of the different sound sources to examine the relative responsiveness of female Mediterranean fruit flies to traps with different acoustic and visual features. The visual attributes of the different sound source assemblies significantly affected capture rates. The range over which the broadcast significantly increased the percentage of female captures was <0.5 m, which may limit the utility of these acoustic cues in large-scale trapping programs. However, the findings of this study do justify further testing of whether optimized short-range acoustic signals could be used to augment longer range pheromonal and visual cues to improve the efficacy of female-selective traps.     

Confusion of Phonotaxis by Masking Sounds in the Bushcricket Conocephalus brevipennis (Tettigoniidae: Conocephalinae)

Female Conocephalus brevipennis, executing phonotaxis to the call of a conspecific male broadcast from a speaker, were subjected to varying levels of background sound so as to progressively mask the attracting signal. The background consisted of either random noise or the call of a sympatric congener, C. nigropleurum and was delivered from above the orienting female. Disruption of phonotaxis was assayed by monitoring the angular dispersion of the female's exit path from the arena, relative to the position of the conspecific call. Random noise significantly interfered with phonotaxis when it was within 2 dB of the conspecific call intensity, both signals measured at the centre of the arena. The equivalent effect of the call C. nigropleurum was 5 dB above conspecific call intensity. The implication of signal loss within background noise is discussed as are the possible interactive behaviours of competing sympatric syntopic species.     

Mosquito bioacoustics: Auditory processing in <Emphasis Type="Italic">Culex pipiens pipiens</Emphasis> L. Males (Diptera,Culicidae) during flight simulation

As a part of mating behavior, male mosquitoes detect and locate females by listening to the sound of their wingbeats. Up to date, the auditory physiological properties in mosquitoes were studied in steady preparations. However, the sensory organs of a flying insect are affected by strong vibrations caused by its own flight muscles and wings. This influence can sufficiently modify the perception due to the nonlinear characteristics of the receptor cells. The aim of this study was to demonstrate the effect of flight conditions on the functioning of Johnston’s organs (hearing organs) in the male mosquitoes Culex pipiens pipiens L. To simulate the flight conditions, a small amount of air around the mosquito was oscillated at 500 Hz along the dorso-ventral axis. These air oscillations affected the pinnate antennae of the mosquito, causing vibration of the antennal flagellum which, in turn, was transmitted to and sensed by Johnston’s organ. Along with the flight simulation, the mosquito was stimulated by low-amplitude sinusoidal auditory signals of different frequencies. The responses were recorded from neurons using glass microelectrodes. The auditory responses contained two rhythms of amplitude modulation which were produced due to nonlinear processes in the receptors: the first rhythm had the difference frequency of flight simulation and the stimulus; the second rhythm corresponded to the beating of heterodyne frequencies. As compared to the steady-state conditions, an additional optimum of auditory sensitivity at 540–640 Hz (the so-called image channel) was shown to appear during flight simulation. This optimum corresponds to the second harmonic of the conspecific female sound. An improvement of sensitivity by 7 dB (2.2-fold) was also observed at the main optimum (220–320 Hz). We conclude that the mechanical influence of locomotor movements not only produces noise but can also improve the sensitivity of the sensory system or even add new properties.     

Spectrum of the calling songs, phonotaxis and the auditory system in the cricket Gryllus bimaculatus]

Behavioural experiments with Y-maze showed that phonotaxis in female crickets to male calling songs (CS) depends on the spectrum of the latter. Conservation of the first low-frequency (5 kc. p. s.) component of the spectrum is the necessary and sufficient condition for the development of normal phonotaxis. Signals which in their temporal characteristics are identical to the CS, but their spectrum contains only high-frequency (12.5 kc. p. s.) component, do no evoke positive phonotaxis. High-frequency signals (10-40 kc. p. s.) induce negative phonotaxis of females in the stationary flight. Beginning from the tympanic organ, the auditory system of crickets exhibits distinct differentiation of elements, which provide the analysis of low- and high-frequency signals. Two types of ascending interneurons transmitting information about the sound from the first auditory center to the brain were described in detail. The first type is associated mainly with low-frequency receptors and effectively transmits all that is necessary for the recognition of temporal characteristics of the CS. The second type presumably accounts for the negative phonotaxis. It is associated mainly with high-frequency receptors, exhibits for the negative phonotaxis. It is associated mainly with high-frequency receptors, exhibits significant after-effect, higher sensitivity to sounds of weak intensities, emphasizes the onset of the stimulus effect, and rapidly habituates to repetitive stimulation.     

Frequency and temporal pattern-dependent phonotaxis of crickets (Teleogryllus oceanicus) during tethered flight and compensated walking

Summary Phonotactic responses ofTeleogryllus oceanicus were studied with two methods. Tethered crickets were stimulated with sound while they performed stationary flight, and steering responses were indicated by abdominal movements. Walking crickets tracked a sound source while their translational movements were compensated by a spherical treadmill, and their walking direction and velocity were recorded.During both flight and walking, crickets attempted to locomote towards the sound source when a song model with 5 kHz carrier frequency was broadcast (positive phonotactic response) and away from the source when a song model with 33 kHz carrier frequency was used (negative phonotactic response) (Figs. 2, 4).One-eared crickets attempted, while flying, to steer towards the side of the remaining ear when stimulated with the 5 kHz model, and away from that side in response to the 33 kHz model (Fig. 3). While walking, one-eared crickets circled towards and away from the intact side in response to the 5 kHz and 33 kHz models, respectively (Fig. 6).Positive and negative responses differed in their temporal pattern requirements. Phonotactic responses were not elicited when a non-calling song pattern (2 pulses/s) was played with a carrier frequency appropriate for positive phonotactic responses (5 kHz), but this pattern did elicit negative responses with 33 kHz carrier frequency (Figs. 7–10). When an intermediate carrier frequency, 15 kHz, was used, the response type (positive or negative) depended on the stimulus temporal pattern; the calling song pattern elicited primarily positive responses, while the non-calling song pattern elicited negative responses (Figs. 11, 12, 14, 15). A curious phenomenon was often observed in the flight steering responses; while most responses to 15 kHz song pattern were primarily positive, they often had an initial negative component which was supplanted by the positive component of the response after approximately 2–5 s (Figs. 11, 12).In recent experiments onGryllus campestris, Thorson et al. (1982) described frequency-dependent errors in phonotactic direction (anomalous phonotaxis) and showed how such errors might arise from the frequency-dependent directional properties of the cricket's auditory apparatus. Our findings, particularly the dependence of response type on temporal pattern when 15 kHz carrier frequency was used, argue that frequency-dependent directional properties alone cannot account for positive and negative phonotaxis inT. oceanicus. Rather, these represent qualitatively different attempts to locomote towards and away from the sound source, respectively.We discuss the possibility that central integration of these opposing tendencies might contribute to anomalous phonotaxis.     

Evidence for short-range sonic communication in lymantriine moths

Sexual communication of nun moth, Lymantria monacha (L.), pink gypsy moth, Lymantria mathura Moore, and fumida tussock moth, Lymantria fumida Butler (all Lepidoptera: Noctuidae: Lymantriinae), is known to be mediated by pheromones. We now show that males are attracted by the sounds of conspecific females over short distances and that wing fanning male and female L. monacha, L. mathura and L. fumida produce species- and sex-specific wing beat and associated click sounds that could contribute to reproductive isolation. Evidence for short-range communication in these lymantriines includes (i) scanning electron micrographs revealing metathoracic tympanate ears, (ii) laser interferometry showing particular sensitivity of tympana tuned to frequency components of sound signals from conspecifics, and (iii) phonotaxis of male L. monacha and L. fumida to speakers playing back sound signals from conspecific females. We conclude that tympanate ears of these moths have evolved in response not only to bat predation, but also for short-range mate finding and possibly recognition.     

The auditory system of blood-sucking mosquito females (Diptera, Culicidae): Acoustic perception during flight simulation

Mosquitoes hear with their plumose antennae which respond to the air movement caused by sound propagation and conduct vibrations to the Johnston’s organ located at the base of each antenna. Each of the two Johnston’s organs contains several tens of thousands mechanosensory cells which detect the displacements of the flagellum and transform them into electric potentials. Hearing plays a very important role in the reproductive behavior of the male mosquitoes. At the same time, our knowledge of hearing in female mosquitoes is very limited and its functional significance is obscure. In this study we measured the auditory sensitivity of female mosquitoes and investigated how the flight conditions affect their hearing. We studied mosquitoes of three species: Anopheles messeae, Aedes excrucians, and Culex pipiens pipiens. The neuronal responses were recorded with a glass microelectrode from the antennal nerve and the deutocerebral interneurons. Stimulation was applied in two modes: (1) the main stimulus against the background of flight simulation (strong vibration with the typical wingbeat frequency of a given mosquito species) and (2) only the main stimulus without the background stimulation. During the flight simulation, females demonstrated an increased sensitivity to frequencies below 200 Hz. The mean auditory receptor threshold at 80–120 Hz was 45 dB, which was 8 dB lower than that without flight simulation. An additional zone of increased sensitivity was also found at frequencies higher than the simulated wingbeat frequency (the so-called image channel). Our analysis of frequency tuning curves measured from the receptors and auditory interneurons shows that mosquito auditory neuronal complex consists of several subsystems which have different frequency tuning parameters, and suggests the possibility of spectral analysis of sounds. Three hypotheses could be proposed on the function of hearing in female mosquitoes: (1) predator avoidance, (2) detection of moving prey, and (3) intraspecific communication. Each of the hypotheses involves the ability to analyze the sound frequency spectrum and subsequent signal recognition.     

Auditory behaviour of a parasitoid fly (Emblemasoma auditrix, Sarcophagidae, Diptera)

Females of the parasitoid fly Emblemasoma auditrix find their host cicada (Okanagana rimosa) by its acoustic signals. In laboratory experiments, fly phonotaxis had a mean threshold of about 66 dB SPL when tested with the cicada calling song. Flies exhibited a frequency dependent phonotaxis when testing to song models with different carrier frequencies (pulses of 6 ms duration and a repetition rate of 80 pulses s(-1)). However, the phonotactic threshold was rather broadly tuned in the range from 5 kHz to 11 kHz. Phonotaxis was also dependent on the temporal parameters of the song models: repetition rates of 60 pulses s(-1) and 80 pulses s and pulse durations of 5-7 ms resulted in the highest percentages of phonotaxis performing animals coupled with the lowest threshold values. Thus, parasitoid phonotaxis is adapted especially to the temporal parameters of the calling song of the host. Choice experiments revealed a preference of a song model with 9 kHz carrier frequency (peak energy of the host song) compared with 5 kHz carrier frequency (electrophysiologically determined best hearing frequency). However, this preference changed with the relative sound pressure level of both signals. When presented simultaneously, E. auditrix preferred 5-kHz signals, if they were 5 dB SPL louder than the 9-kHz signal.     

Phonotaxis to male’s calls embedded within a chorus by female gray treefrogs, Hyla versicolor

During the reproductive season, male Hyla versicolor produce advertisement calls to attract females. Females exhibit phonotaxis and approach the individual callers, resulting in amplexus. For frogs that call from dense choruses, the extent to which and the range from which a male’s advertisement call within a chorus can be heard by a receptive female leading to phonotaxis is unclear. We investigated females’ responses to natural choruses in the field and found that they were attracted and showed directed orientation to breeding choruses at distances up to 100 m. To assess the role of acoustic cues in the directed orientation, we conducted acoustic playback experiments in the laboratory using conspecific call and noise as stimuli, as well as chorus sounds (that contained calls from a focal male) recorded at various distances, all played at naturalistic intensities. Using two response metrics (females’ normalized response times and their phonotaxis trajectories) we found that, unlike the field experiments, females oriented and were attracted to chorus sounds from 1 to 32 m only, but not from >32 m, or to band-limited noise. Possible reasons for the observed difference in phonotaxis behavior in the two experimental conditions were discussed.     

Acoustic irradiation produced by flying moths (Lepidoptera,Noctuidae)

Characteristics of acoustic waves accompanying the flight of noctuid moths (Noctuidae) were measured. The low-frequency part of the spectrum is formed of a series of up to 17 harmonics of the wingbeat frequency (30–50 Hz) with a general tendency toward the decrease in the spectral density and the increase in the sound frequency. The root-mean-square level of the sound pressure from flapping wings was found to be 70–78 dB SPL. Besides low-frequency components, the flight of moths was accompanied by short ultrasonic pulses, which appeared with every wingbeat. Most of the spectral energy was concentrated within a range of 7–150 kHz with the main peaks at 60–110 kHz. The short-term pulses were divided into two or more subpulses with different spectra. The high-frequency pulses were produced at two phases of the wingbeat cycle: during the pronation of the wings at the highest point and at the beginning of their upward movement from the lowest point. In most of the specimens tested, the peak amplitude of sounds varied from 55 to 65 dB SPL at a distance of 6 cm from the insect body. However, in nine noctuid species, no high-frequency acoustic components were recorded. In these experiments, the acoustic flow from the flying moth within a frequency range of 2 to 20 kHz did not exceed the self-noise level of the microphone amplifier (RMS 18 dB SPL). Probable mechanisms of the high frequency acoustic emission during flight, the effect of these sounds on the auditory sensitivity of moths, and the possibility of their self-revealing to insectivorous bats are discussed. In addition, spectral characteristics of the moth echolocation clicks were more precisely determined within the higher frequency range (>100 kHz).     

Encoding properties of auditory neurons in the brain of a soniferous damselfish: response to simple tones and complex conspecific signals

The fish auditory system encodes important acoustic stimuli used in social communication, but few studies have examined response properties of central auditory neurons to natural signals. We determined the features and responses of single hindbrain and midbrain auditory neurons to tone bursts and playbacks of conspecific sounds in the soniferous damselfish, Abudefduf abdominalis. Most auditory neurons were either silent or had slow irregular resting discharge rates <20 spikes s⁻¹. Average best frequency for neurons to tone stimuli was ~130 Hz but ranged from 80 to 400 Hz with strong phase-locking. This low-frequency sensitivity matches the frequency band of natural sounds. Auditory neurons were also modulated by playbacks of conspecific sounds with thresholds similar to 100 Hz tones, but these thresholds were lower than that of tones at other test frequencies. Thresholds of neurons to natural sounds were lower in the midbrain than the hindbrain. This is the first study to compare response properties of auditory neurons to both simple tones and complex stimuli in the brain of a recently derived soniferous perciform that lacks accessory auditory structures. These data demonstrate that the auditory fish brain is most sensitive to the frequency and temporal components of natural pulsed sounds that provide important signals for conspecific communication.     

Male responses to conspecific advertisement signals in the field cricket Gryllus rubens (Orthoptera: Gryllidae)

In many species males aggregate and produce long-range advertisement signals to attract conspecific females. The majority of the receivers of these signals are probably other males most of the time, and male responses to competitors' signals can structure the spatial and temporal organization of the breeding aggregation and affect male mating tactics. I quantified male responses to a conspecific advertisement stimulus repeatedly over three age classes in Gryllus rubens (Orthoptera: Gryllidae) in order to estimate the type and frequency of male responses to the broadcast stimulus and to determine the factors affecting them. Factors tested included body size, wing dimorphism, age, and intensity of the broadcast stimulus. Overall, males employed acoustic response more often than positive phonotactic response. As males aged, the frequency of positive phonotactic response decreased but that of the acoustic response increased. That is, males may use positive phonotaxis in the early stages of their adult lives, possibly to find suitable calling sites or parasitize calling males, and then later in life switch to acoustic responses in response to conspecific advertisement signals. Males with smaller body size more frequently exhibited acoustic responses. This study suggests that individual variation, more than any factors measured, is critical for age-dependent male responses to conspecific advertisement signals.     

雌蚊翅振音及其在蚊虫防治中的应用

蚊虫飞翔时 ,翅上下拍打会形成连续的翅振音。雌蚊翅振音频率会随蚊种、蚊体长和日龄及环境温度而变化 ,一般在 3 0 0～ 5 0 0Hz之间。同种个体间翅振音频率变化较小 ,常在平均频率± 5 0Hz范围内。雄蚊只对基本频率的雌蚊翅振音起反应 ,雌蚊翅振音的偶然变化不会降低其对雄蚊的引诱力。雄蚊对雌蚊翅振音具有敏感反应的主要原因是其听觉器官对雌蚊翅振音形成了良好的适应性。因此 ,许多蚊虫研究者希望利用雌蚊翅振音来防治蚊虫。因该方法对环境安全 ,在今后蚊虫防治中其重要性将会日益显现。     

Fluttering wing feathers produce the flight sounds of male streamertail hummingbirds

Sounds produced continuously during flight potentially play important roles in avian communication, but the mechanisms underlying these sounds have received little attention. Adult male Red-billed Streamertail hummingbirds (Trochilus polytmus) bear elongated tail streamers and produce a distinctive 'whirring' flight sound, whereas subadult males and females do not. The production of this sound, which is a pulsed tone with a mean frequency of 858 Hz, has been attributed to these distinctive tail streamers. However, tail-less streamertails can still produce the flight sound. Three lines of evidence implicate the wings instead. First, it is pulsed in synchrony with the 29 Hz wingbeat frequency. Second, a high-speed video showed that primary feather eight (P8) bends during each downstroke, creating a gap between P8 and primary feather nine (P9). Manipulating either P8 or P9 reduced the production of the flight sound. Third, laboratory experiments indicated that both P8 and P9 can produce tones over a range of 700-900 Hz. The wings therefore produce the distinctive flight sound, enabled via subtle morphological changes to the structure of P8 and P9.     

ACOUSTIC COMMUNICATION AND MATING BEHAVIOUR IN WATER BUGS OF THE GENUS MICRONECTA

ABSTRACT

The role of male sounds in attracting females and in mating was investigated in the three most common species of Micronecta found in ponds 60 km NE of Melbourne, Australia: M. concordia, M. tasmanica and M. robusta. In playback experiments using recorded male signals, females were attracted to signals of conspecific males, in preference to signals of heterospecific sympatric males. Studies of mating behaviour, using video recording, showed that signals were obligatory for mating. These findings strongly suggest that acoustic signalling is important in reproductive isolation in Micronecta. Comparisons between matings and rejected matings showed that, within each species, copulation only occurred following a certain minimum number of acoustic signals with more pulse-trains than the mean for the species. Low amplitude sounds (after signals) were also important in courtship, immediately preceding mating. No sounds occurred during copulation.     

Directional hearing in a silicon cricket

Phonotaxis is the ability to orient towards or away from sound sources. Crickets can locate conspecifics by phonotaxis to the calling (mating) song they produce, and can evade bats by negative phonotaxis from echolocation calls. The behaviour and underlying physiology have been studied in some depth, and the auditory system solves this complex problem in a unique manner. Experiments conducted on a simulation model of the system indicated that the mechanism output a directional signal to sounds ahead at calling song frequency and to sounds behind at echolocation frequencies. We suggest that this combination of responses helps simplify later processing in the cricket. To further explore this result, an analogue, very large scale integrated (aVLSI) circuit model of the mechanism was designed and built; results from testing this agreed with the simulation. The aVLSI circuit was used to test a further hypothesis about the potential advantages of the positioning of the acoustic inputs for sound localisation during walking. There was no clear advantage to the directionality of the system in their location. The aVLSI circuitry is now being extended to use on a robot along with previously modelled neural circuitry to better understand the complete sensorimotor pathway.