Low-Frequency Sounds Repel Male Mosquitoes <Emphasis Type="Italic">Aedes diantaeus</Emphasis> N.D.K. (Diptera,Culicidae)

We experimentally demonstrated that tonal acoustic signals with a carrier frequency of 140–200 Hz had a repellent effect on male mosquitoes (Culicidae). Swarming males of Aedes diantaeus were concentrated in a small space near the auxiliary attracting sound source which simulated the flight sound of conspecific females (carrier frequency 280–320 Hz). Then, the resulting cluster of attracted mosquitoes was stimulated with test signals of variable amplitude and carrier frequency from a second loudspeaker. The direction of mosquito flight from the source of test sounds and a decrease in their number above the attracting sound source were used as the criteria of behavioral response. Pronounced avoidance responses (negative phonotaxis) of swarming mosquitoes were observed in the range of 140–200 Hz. Most of the mosquitoes left the area above the attracting sound source within one second after the onset of the test signal. Mosquitoes mostly flew up, sideways, and backwards in relation to the test acoustic vector. We presume that mosquitoes develop defensive behavior against attacking predatory insects based on analysis of auditory information. The range of negative phonotaxis is limited at higher frequencies by the spectrum of the flight sounds of conspecific females, and in the low frequency range, by the increasing level of atmospheric noise.     

Avoidance Responses to Infrasound in Downstream Migrating European Silver Eels, Anguilla anguilla

In an attempt to develop an efficient acoustic fish fence, we have designed an infrasound source able to generate large nearfield particle acceleration. The source generates water movements by means of two symmetrical pistons in an air-filled cylinder with 21cm bore. The pistons are driven by eccentric coupling to an electric motor, with 5cm p.p. amplitude. The piston movements are 180° out of phase. The piston reaction forces are thus opposed, leading to vibration free operation. The submergible infrasound source is operated freely suspended in the water mass. The emitted sound frequency is 11.8Hz. The particle acceleration is about 0.01ms^–2 at a distance of 3m, corresponding to the threshold intensity for deterring effects of infrasound on Atlantic salmon smolts. The sound source was employed to test the effect of intense infrasound on migrating European silver eels. Fish confined in a tank displayed startle behaviour and prolonged stress reactions, telemetrically monitored as tachycardia, in response to intense infrasound. The field tests were carried out in the River Imsa. A trap that catches all the descending eels is installed near the river mouth. The trap was separated in four equal sections. During the periods with infrasound exposure, the proportion of silver eels entering the section closest to the sound source was reduced to 43% of the control value. In the section closest to the opposite river bank, infrasound increased the proportion of trapped eels to 144% of the control values. This shift of the migrating eels away from the infrasound source was highly significant.     

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.     

Ultrasonic startle behavior in bushcrickets (Orthoptera; Tettigoniidae)

1. In the present work, we show that in flight, bushcrickets not previously known to respond to ultrasound alter their flight course in response to ultrasonic stimuli. Such stimuli elicit in flying Neoconocephalus ensiger an extension of the front and middle legs along the body and a rapid closure of all 4 wings (Fig. 1). This is a short latency acoustic startle response to ultrasound, consistent with acoustic startle responses of other insects. 2. The percentage of trials on which acoustic startle responses were elicited was maximum (90%) for sound frequencies ranging from 25 to at least 60 kHz. No acoustic startle response was observed at frequencies of 5 or 10 kHz (Fig. 2). The threshold for the response was roughly 76 dB between 25 to 60 kHz (Fig. 2) and the behavioral latency was 45 ms (Fig. 3). Recordings from flight muscles show that they cease discharging during the acoustic startle response (Fig. 4). 3. The characteristics of the acoustic startle response match those of an auditory interneuron called the T-neuron. The frequency sensitivity of this neuron is greatest for sound frequencies ranging from 13 to 60 kHz (Fig. 6). Moreover, we found that the neuron produces many more spikes to ultrasound (30 kHz) of increasing intensities than to a conspecific communication sound, whose dominant frequency is 14 kHz (Fig. 7).     

Common sole larvae survive high levels of pile-driving sound in controlled exposure experiments

In view of the rapid extension of offshore wind farms, there is an urgent need to improve our knowledge on possible adverse effects of underwater sound generated by pile-driving. Mortality and injuries have been observed in fish exposed to loud impulse sounds, but knowledge on the sound levels at which (sub-)lethal effects occur is limited for juvenile and adult fish, and virtually non-existent for fish eggs and larvae. A device was developed in which fish larvae can be exposed to underwater sound. It consists of a rigid-walled cylindrical chamber driven by an electro-dynamical sound projector. Samples of up to 100 larvae can be exposed simultaneously to a homogeneously distributed sound pressure and particle velocity field. Recorded pile-driving sounds could be reproduced accurately in the frequency range between 50 and 1000 Hz, at zero to peak pressure levels up to 210 dB re 1μPa(2) (zero to peak pressures up to 32 kPa) and single pulse sound exposure levels up to 186 dB re 1μPa(2)s. The device was used to examine lethal effects of sound exposure in common sole (Solea solea) larvae. Different developmental stages were exposed to various levels and durations of pile-driving sound. The highest cumulative sound exposure level applied was 206 dB re 1μPa(2)s, which corresponds to 100 strikes at a distance of 100 m from a typical North Sea pile-driving site. The results showed no statistically significant differences in mortality between exposure and control groups at sound exposure levels which were well above the US interim criteria for non-auditory tissue damage in fish. Although our findings cannot be extrapolated to fish larvae in general, as interspecific differences in vulnerability to sound exposure may occur, they do indicate that previous assumptions and criteria may need to be revised.     

Comparison of lung sound transducers using a bioacoustic transducer testing system.

Sensors used for lung sound research are generally designed by the investigators or adapted from devices used in related fields. Their relative characteristics have never been defined. We employed an artificial chest wall with a viscoelastic surface and a white noise signal generator as a stable source of sound to compare the frequency response and pulse waveform reproduction of a selection of devices used for lung sound research. We used spectral estimation techniques to determine frequency response and cross-correlation of pulses to determine pulse shape fidelity. The sensors evaluated were the Siemens EMT 25 C accelerometer (Siemens); PPG 201 accelerometer (PPG); Sony ECM-T150 electret condenser microphone with air coupler (air coupler; with cylindrical air chambers of 5-, 10-, and 15-mm diameter and conical air chamber of 10-mm diameter); Littman classic stethoscope head (Littman) connected to an electret condenser microphone; and the Andries Tek (Andries) electronic stethoscope. We found that the size and shape of the air coupler chamber to have no important effect on the detected sound. The Siemens, air coupler, and Littman performed similarly with relatively flat frequency responses from 200 to 1,200 Hz. The PPG had the broadest frequency response, with useful sensitivity extending to 4,000 Hz. The Andries' frequency response was the poorest above 1,000 Hz. Accuracy in reproducing pulses roughly corresponded with the high-frequency sensitivity of the sensors. We conclude that there are important differences among commonly used lung sound sensors that have to be defined to allow the comparison of data from different laboratories.     

Sound localization by the Hawaiian squirrelfishes, Myripristis berndti and M. argyromus

Sound localization was investigated in a large pond open to a bay and similar to the normal environment of the animals. Observations were made of fish movements towards one of two underwater loud-speakers emitting squirrelfish alarm calls normally produced in response to predators. When the sound source was within 2·0 m of the test cage housing the fish, the subjects faced and moved toward the speaker. The animals responded some of the time when the source was within 3·0 m but generally did not orient to the sound source when the speaker was beyond 3·0 m. Response loss was correlated with the fish being in the acoustic far-field. Possible cues which release and direct localization remain unknown, but include particle velocity information alone, or some change in particle velocity: pressure relationships.     

Modulation of auditory responsiveness in the locust

The auditory responsiveness of a number of neurones in the meso- and metathoracic ganglia of the locust, Locusta migratoria, was found to change systematically during concomitant wind stimulation. Changes in responsiveness were of three kinds: a suppression of the response to low frequency sound (5 kHz), but an unchanged or increased response to high frequency (12 kHz) sound; an increased response to all sound; a decrease in the excitatory, and an increase in the inhibitory, components of a response to sound. Suppression of the response to low frequency sound was mediated by wind, rather than by the flight motor. Wind stimulation caused an increase in membrane conductance and concomitant depolarization in recorded neurones. Wind stimulation potentiated the spike response to a given depolarizing current, and the spike response to a high frequency sound, by about the same amount. The strongest wind-related input to interneuron 714 was via the metathoracic N6, which carries the axons of auditory receptors from the ear. The EPSP evoked in central neurones by electrical stimulation of metathoracic N6 was suppressed by wind stimulation, and by low frequency (5 kHz), but not high frequency (10 kHz), sound. This suppression disappeared when N6 was cut distally to the stimulating electrodes. Responses to low frequency (5 kHz), rather than high frequency (12 kHz), sounds could be suppressed by a second low frequency tone with an intensity above 50-55 dB SPL for a 5 kHz suppressing tone. Suppression of the electrically-evoked EPSP in neurone 714 was greatest at those sound frequencies represented maximally in the spectrum of the locust's wingbeat. It is concluded that the acoustic components of a wind stimulus are able to mediate both inhibition and excitation in the auditory pathway. By suppressing the responses to low frequency sounds, wind stimulation would effectively shift the frequency-response characteristics of central auditory neurones during flight.     

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.     

Range of corixid sound signals in the biotope

ABSTRACT. The propagation of sound in the frequency band (2–12 kHz) used by the Corixidae was measured in two shallow natural ponds. At distances of more than 1 m from the shore the water was at least O.4 m deep. The first pond was eutrophic and contained no plants. The spread of sound into the open water obeyed approximately the geometric attenuation of the sound pressure level (SPL); a loss of 6dB for each doubling of the distance from the point sound source. Near the shore the attenuation was considerably greater, especially for low frequencies; for a 2 kHz signal the damping of the SPL was c. 40–50 dB/m.
The second pond had dense plant growth, and the sound attenuation depended strongly on the photosynthetic activity of the waterplants. Measurements in winter, with an overcast sky, revealed only a slight damping effect of the plants for a 10kHz test signal. During intense sunlight in summer, however, in addition to the geometric attenuation the damping effect of the plants over a distance of O.5 m was 50 dB for a 2 kHz signal and 80 dB for 10 kHz. This effect was due to gas bubbles produced during intense photosynthesis.
Song A of Corixa dentipes Thms. (Heteroptera) males elicits usually a response by male conspecifics. The threshold SPL for this response was measured to be c. 40 dB lower than the SPL at a distance of O.1 m from a stridulating animal. From the measurements of sound propagation it follows, therefore, that the effective range of Song A in the most favourable case is at least 10m, though in a pond overgrown with plants it can be less than O.4 m.     

Awareness reactions and avoidance responses to sound in juvenile Atlantic salmon, Salmo salar L.

The possibility of using intense sound as a deterrent for juvenile Atlantic salmon ( Sulmo salar L. ) was studied by recording both physiological awareness reactions in an acoustic tube and behavioural avoidance responses in a pool. The measured awareness reactions consisted of decreased heart rate and breathing movements. Three criteria for the awareness reaction were used to compare the effect of different frequencies between 5 and 150 Hz: (i) threshold for spontaneous awareness reactions relative to the hearing thresholds, (ii) magnitude of the change in heart rate, (iii) degree of habituation to sound. After these criteria the lowest frequencies (5–10 Hz) were most effective in eliciting an awareness reaction from the fish. The responses of freely swimming fish to 10 and 150 Hz sounds were studied in an artificial pool. Juvenile salmon showed avoidance responses to 10 Hz stimulation at intensities 10–15 dB above the threshold for spontaneous awareness reactions measured in the acoustic tube. The 150 Hz sound failed to evoke avoidance responses, even at a level 30 dB above the threshold for spontaneous awareness reactions.     

Extinction of a conditioned response in rainbow trout selected for high or low responsiveness to stress

Two lines of rainbow trout (Oncorhynchus mykiss) that exhibit divergent endocrine responsiveness to stressors also display disparate behavioral traits. To investigate whether the high-responding (HR) and low-responding (LR) fish also differ in cognitive function, the rate of extinction of a conditioned response was compared between the two lines. Groups of HR and LR fish were exposed to a paired conditioned stimulus (CS; water off) and unconditioned stimulus (US; confinement stressor). After exposure to 18 CS-US pairings, at least 70% of individuals of both lines acquired a conditioned response (CR) manifested as an elevation of blood cortisol levels on presentation of the CS only. Post-conditioning, the fish were tested by presentation of the CS at weekly intervals, for 4 weeks, with no further reinforcement, and the extinction of the CR in the two lines was compared. The decline in mean plasma cortisol levels after exposure to the CS over successive tests suggested that the CR was retained for a shorter period among the HR (<14 days) than LR fish (<21 days). The frequency of individuals within each line whose plasma cortisol levels indicated a stress response when exposed to the CS was significantly greater among the LR than HR fish at 14 and 21 days with no HR fish falling into this category at 21 days. At 28 days post-conditioning, there were no HR fish and only three LR fish were categorized as "stressed". These results suggest that there are differences in cognitive function between the two lines. Possible mechanisms underlying these differences are discussed.     

Directional orientation of pomacentrid larvae to ambient reef sound

The mechanisms by which reef fish larvae locate settlement habitat at the end of their pelagic phase are unclear. We used an in situ binary choice chamber and an artificial source of reef sound to determine whether pomacentrid larvae can use ambient sound to locate reefs. Larvae were caught in light traps and then placed in a submerged binary choice chamber with an artificial source of reef sound ~80 m from one end of the chamber. At night, larvae moved towards the sound source; during the day, larvae showed no preference. These results suggest that pomacentrid larvae can detect reef sound and are capable of directional hearing. While other studies have shown that reef fish larvae respond to reef sound, and that the adults of some species can localize underwater sound sources, the localization of underwater sound by fish larvae has not been demonstrated previously.Communicated by Ecological Editor P.F. Sale     

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。     

Field and laboratory electroantennographic measurements of pheromone plume structure correlated with oriental fruit moth behaviour

Abstract. Peak-to-trough electroantennogram amplitudes (bursts), caused by the individual filaments of a plume of female pheromone, diminish as high-emission-rate sources are approached by male Grapholita molesta , and this reduction is correlated with in-flight arrestment (ceasing to advance upwind). These findings are consistent with the hypothesis that one cause of in-flight arrestment in response to high-concentration point sources is the attenuation of the peak-to- trough amplitudes close to the source. High burst frequency, high pheromone flux, or low levels of continuous neuronal activity all are less well correlated with arrestment. Rather, arrestment appears due to a reduction of chemosensory input to the CNS during flight up the plume, even though the actual molecular concentration continues to increase. In a laboratory wind tunnel, upwind flight initiation by more than 20% of males was elicited only by pheromone source concentrations evoking significant fluctuations in EAG amplitudes at downwind release points. The burst frequencies that evoked high levels of upwind flight initiation ranged from a mean of 0.4-2.2 bursts/s. Because a previous study revealed that flying male G. molesta change their course angle within 0.15 s of losing or contacting pheromone, these EAG burst frequencies indicate that during flight in a pheromone plume, many manoeuvres are probably made in response to contact with individual plume filaments. Thus, upwind flight tracks may be shaped by hundreds of steering reactions in response to encounters with individual pheromone filaments and pockets of clean air. Field-recorded EAGs reveal that burst amplitudes diminish from 3 to 30 m downwind of the source, whereas burst frequencies do not, averaging c. 1/s at 3, 10 and 30 m downwind.     

3-D-orientation with the octavolateralis system.

Fish detect and localize a sound source with inner ear receptors and with the mechanosensory lateral line. The inner ear of fish is sensitive to the water displacements caused by sound waves through a direct, inertial response by hair cell epithelia of the ear. Hearing specialists, such as goldfish and herring, have accessory peripheral structures that provide additional sensitivity to the pressure component of a sound wave. While the inner ear of fish responds to the whole body motions caused by sound waves and--in case of hearing specialists--to sound pressure, the lateral line is only sensitive to water motions relative to the surface of the fish and to local pressure gradients. Using lateral line and/or acoustic input, some fish can determine the direction and the distance to a sound source. Most likely they do so by exploiting some of the mechanisms described in this paper. Piscivorous fish may use lateral line input to detect the wakes caused by swimming fish. Even in the absence of light catfish, for instance, can follow a 10 s old, three-dimensional wake left by a prey fish over distances up to 55 prey-body length.     

UNDERWATER NOISE OF WHALE-WATCHING BOATS AND POTENTIAL EFFECTS ON KILLER WHALES (ORCINUS ORCA), BASED ON AN ACOUSTIC IMPACT MODEL

Underwater noise of whale-watching boats was recorded in the popular killer whale-watching region of southern British Columbia and northwestern Washington State. A software sound propagation and impact assessment model was applied to estimate zones around whale-watching boats where boat noise was audible to killer whales, where it interfered with their communication, where it caused behavioral avoidance, and where it possibly caused hearing loss. Boat source levels ranged from 145 to 169 dB re 1 μPa @ 1 m, increasing with speed. The noise of fast boats was modeled to be audible to killer whales over 16 km, to mask killer whale calls over 14 km, to elicit a behavioral response over 200 m, and to cause a temporary threshold shift (TTS) in hearing of 5 dB after 30–50 min within 450 m. For boats cruising at slow speeds, the predicted ranges were 1 km for audibility and masking, 50 m for behavioral responses, and 20 m for TTS. Superposed noise levels of a number of boats circulating around or following the whales were close to the critical level assumed to cause a permanent hearing loss over prolonged exposure. These data should be useful in developing whale-watching regulations. This study also gave lower estimates of killer whale call source levels of 105–124 dB re 1 μPa.     

Aggressive behaviour of underyearling rainbow trout in simulated winter concealment habitat

The behaviour of wild underyearling rainbow trout Oncorhynchus mykiss in concealment habitat in a laboratory stream aquarium at 2 and 6°C was recorded daily with an infrared video camera for 90 min over dawn. Aggressive behaviours (threat nips, nips and chases) were frequent during this time as fish entered concealment habitat. Aggressiveness varied widely among fish groups, with a range of 1 to 45 aggressive acts being initiated during a 90-min filming period. Larger fish initiated most of the encounters and removed a higher proportion of fish from concealment than did smaller aggressors. Thirteen per cent of the aggressive acts resulted in the recipient being completely removed from concealment habitat. The highest combined frequency of aggressive acts was 0.64 per visible fish per 10 min period and occurred at relatively low light levels (300 lx). The data suggest that when fish density is high, such as when habitat is limited, aggressive behaviour in winter may cause some underyearling salmonids to be excluded from concealment.     

Swarming behaviour of the mosquito Culex pipiens quinquefasciatus: a quantitative analysis

ABSTRACT. Analysis of video recordings of Culex pipiens quinquefasciatus Say swarms showed that the overall swarming patterns of males and virgin females are similar, even though the short-term characteristics of their flight, such as speed and turning angle, differ. This suggests they have the same response to the visual cue of the swarm 'marker'. The swarming pattern of an individual consists of elliptical loops which, in the short-term, define an area smaller than the swarm as a whole. The foci of these ellipses drift gradually with respect to the marker; individuals do not seem to have preferred positions within the swarm, but drift at random. Male mosquitoes identify and locate females by the sound of their wing-beats. Diffuse sound at the wing-beat frequency of female C.p. quinquefasciatus (500–600 Hz) caused males within the swarm to slow down significantly. Their turning behaviour remained unchanged, so the ellipses decreased in size. The swarm as a whole collapsed into a smaller volume, centred over the marker. Apparently, the response of males to sound consists of at least two parts: initially they alter their flight speed, and then alter their turning behaviour once they have located the source of the sound.     

Ontogeny of Acoustic and Feeding Behaviour in the Grey Gurnard, Eutrigla gurnardus

Although sound production in teleost fish is often associated with territorial behaviour, little is known of fish acoustic behaviour in other agonistic contexts such as competitive feeding and how it changes during ontogeny. The grey gurnard, Eutrigla gurnardus, frequently emits knock and grunt sounds during competitive feeding and seems to adopt both contest and scramble tactics under defensible resource conditions. Here we examine, for the first time, the effect of fish size on sound production and agonistic behaviour during competitive feeding. We have made sound (alone) and video (synchronized image and sound) recordings of grey gurnards during competitive feeding interactions. Experimental fish ranged from small juveniles to large adults and were grouped in four size classes: 10–15, 15–20, 25–30 and 30–40 cm in total length. We show that, in this species, both sound production and feeding behaviour change with fish size. Sound production rate decreased in larger fish. Sound duration, pulse duration and the number of pulses increased whereas the peak frequency decreased with fish size, in both sound types (knocks and grunts). Interaction rate and the frequency of agonistic behaviour decreased with increasing fish size during competitive feeding sessions. The proportion of feeding interactions accompanied by sound production was similar in all size classes. However, the proportion of interactions accompanied by knocks (less aggressive sounds) and by grunts (more aggressive) increased and decreased with fish size, respectively. Taken together, these results suggest that smaller grey gurnards compete for food by contest tactics whereas larger specimens predominantly scramble for food, probably because body size gives an advantage in locating, capturing and handling prey. We further suggest that sounds emitted during feeding may potentially give information on the motivation and ability of the individual to compete for food resources.