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.     

Sounds audible to migrating birds

We have measured sound levels from frog choruses in eastern New York State at altitudes of up to several hundred metres. Rana pipiens choruses from small ponds often could be recorded by a radio microphone up to 500 m, and on an especially favourable night with light winds they were clearly audible even at 965 m at about 20 dB SPL in the 1·5 to 2·5 2·5 kHz frequency band. Sound travels upward much farther and more predictably than along the surface. Many natural sounds, including those from frogs, insects, whitecaps, and perhaps wind-blown vegetation, arise from large areas and therefore act as extended sources. The intensities of such sounds decrease with altitude more slowly than expected from the inverse square law. Natural sound fields provide migrating birds with a potential source of information about the kind of land or water below them, and their progress over acoustic landmarks could inform them about wind velocity. Because atmospheric absorption increases with frequency, several hundred metres of air act as a low-pass filter, so that altitude could be estimated from the relative reduction of higher frequencies in a familiar sound.     

Cartilage Conduction Is Characterized by Vibrations of the Cartilaginous Portion of the Ear Canal

Cartilage conduction (CC) is a new form of sound transmission which is induced by a transducer being placed on the aural cartilage. Although the conventional forms of sound transmission to the cochlea are classified into air or bone conduction (AC or BC), previous study demonstrates that CC is not classified into AC or BC (Laryngoscope 124: 1214–1219). Next interesting issue is whether CC is a hybrid of AC and BC. Seven volunteers with normal hearing participated in this experiment. The threshold-shifts by water injection in the ear canal were measured. AC, BC, and CC thresholds at 0.5–4 kHz were measured in the 0%-, 40%-, and 80%-water injection conditions. In addition, CC thresholds were also measured for the 20%-, 60%-, 100%-, and overflowing-water injection conditions. The contributions of the vibrations of the cartilaginous portion were evaluated by the threshold-shifts. For AC and BC, the threshold-shifts by the water injection were 22.6–53.3 dB and within 14.9 dB at the frequency of 0.5–4 kHz, respectively. For CC, when the water was filled within the bony portion, the thresholds were elevated to the same degree as AC. When the water was additionally injected to reach the cartilaginous portion, the thresholds at 0.5 and 1 kHz dramatically decreased by 27.4 and 27.5 dB, respectively. In addition, despite blocking AC by the injected water, the CC thresholds in force level were remarkably lower than those for BC. The vibration of the cartilaginous portion contributes to the sound transmission, particularly in the low frequency range. Although the airborne sound is radiated into the ear canal in both BC and CC, the mechanism underlying its generation is different between them. CC generates airborne sound in the canal more efficiently than BC. The current findings suggest that CC is not a hybrid of AC and BC.     

Biophysics of underwater hearing in the clawed frog,Xenopus laevis

Anesthetized clawed frogs (Xenopus laevis) were stimulated with underwater sound and the tympanic disk vibrations were studied using laser vibrometry. The tympanic disk velocities ranged from 0.01 to 0.5 mm/s (at a sound pressure of 2 Pa) in the frequency range of 0.4–4 kHz and were 20–40 dB higher than those of the surrounding tissue. The frequency response of the disk had two peaks, in the range of 0.6–1.1 kHz and 1.6–2.2 kHz, respectively. The first peak corresponded to the peak vibrations of the body wall overlying the lung. The second peak matched model predictions of the pulsations of the air bubble in the middle ear cavity. Filling the middle ear cavity with water lowered the disk vibrations by 10–30 dB in the frequency range of 0.5–3 kHz.Inflating the lungs shifted the low-frequency peak downwards, but did not change the high-frequency peak. Thus, the disk vibrations in the frequency range of the mating call (main energy at 1.7–1.9 kHz) were mainly caused by pulsations of the air in the middle ear cavity; sound transmission via the lungs was more important at low frequencies (below 1 kHz). Furthermore, the low-frequency peak could be reversibly reduced in amplitude by loading the larynx with metal or tissue glue. This shows that the sound-induced vibrations of the lungs are probably coupled to the middle ear cavities via the larynx. Also, anatomical observations show that the two middle ear cavities and the larynx are connected in an air-filled recess in submerged animals.This arrangement is unique to pipid frogs and may be a structural adaptation to connect all the air spaces of the frog and improve low-frequency underwater hearing. Another function of the recess may be to allow cross-talk between the two middle ear cavities. Thus, the ear might be directional. Our pilot experiments show up to 10 dB difference between ipsi- and contralateral stimulus directions in a narrow frequency range around 2 kHz.     

Small tympanic membrane perforations in the inferior quadrants do not impact the manubrium vibration in guinea pigs

Background

It has been believed that location of the perforation has a significant impact on hearing loss. However, recent studies have demonstrated that the perforation sites had no impact on hearing loss. We measured the velocity and pattern of the manubrium vibration in guinea pigs with intact and perforated eardrum using a laser Doppler vibrometer in order to determine the effects of different location perforations on the middle ear transfer functions.

Methods

Two bullas from 2 guinea pigs were used to determine stability of the umbo velocities, and 12 bullas from six guinea pigs to determine the effects of different location perforations on sound transmission. The manubrium velocity was measured at three points on the manubrium in the frequencies of 0.5–8 kHz before and after a perforation was made. The sites of perforations were in anterior-inferior (AI) quadrants of left ears and posterior-inferior (PI) quadrants of right ears.

Results

The manubrium vibration velocity losses were noticed in the perforated ears only below 1.5 kHz. The maximum velocity loss was about 7 dB at 500 Hz with the PI perforation. No significant difference in the velocity loss was found between AI and PI perforations. The average ratio of short process velocity to the umbo velocity was approximately 0.5 at all frequencies. No significant differences were found before and after perforation at all frequencies (p>0.05) except 7 kHz (p = 0.004) for both AI and PI perforations.

Conclusions

The manubrium vibration velocity losses from eardrum perforation were frequency-dependent and the largest losses occur at low frequencies. Manubrium velocity losses caused by small acute inferior perforations in guinea pigs have no significant impact on middle ear sound transmission at any frequency tested. The manubrium vibration axis may be perpendicular to the manubrium below 8 kHz in guinea pigs.     

The acoustic behaviour of the bushcricket Tettigonia cantans II. Transmission of airborne-sound and vibration signals in the biotope

The airborne-sound and the vibratory signals produced by stridulating $\text{Tettigonia cantans}$ males, and the transmission of these signals in the natural biotope were investigated.The song of $\text{T. cantans}$ is composed of repeated uniform syllables with a rate of ca. 30/sec. Intensity approaches 100 dB SPL, 10 cm away from the animal. The spectrum shows three dominant frequency ranges around 8, 16 and 32 kHz.Airborne transmission of the song in such vegetation layers as are found in the biotopes of $\text{T. cantans}$ shows an excess attenuation which increases with frequency. The relative intensities of the frequency components of the song vary as a result of the kind of vegetation, the positions of emitter and receiver, and the separation distance. These relative differences in intensity may be useful during the phonotactic approach to conspecific partners, providing a measure of the distance from the sound source.Stridulating males also produce vibratory signals in the plants they sit on. The spectrum of these signals includes frequencies up to 8 kHz, the first dominant frequency of the song: low frequency components are induced in the plants via the legs and abdomen of the animal. The vibratory signals are transmitted mainly in the form of bending waves. Near the animal, amplitude modulation corresponds to that of the song. At greater distances, reflections and frequency-dependent propagation velocities, cause distortions of this time pattern. Transmission depends greatly on the mechanical properties of the particular plant, attenuation values of 20–50 dB/m being found. Nevertheless, in most cases, vibratory signals may be perceived up to 1.5 – 2 metres away from a stridulating male.     

Chronic exposure to low frequency noise at moderate levels causes impaired balance in mice

We are routinely exposed to low frequency noise (LFN; below 0.5 kHz) at moderate levels of 60-70 dB sound pressure level (SPL) generated from various sources in occupational and daily environments. LFN has been reported to affect balance in humans. However, there is limited information about the influence of chronic exposure to LFN at moderate levels for balance. In this study, we investigated whether chronic exposure to LFN at a moderate level of 70 dB SPL affects the vestibule, which is one of the organs responsible for balance in mice. Wild-type ICR mice were exposed for 1 month to LFN (0.1 kHz) and high frequency noise (HFN; 16 kHz) at 70 dB SPL at a distance of approximately 10-20 cm. Behavior analyses including rotarod, beam-crossing and footprint analyses showed impairments of balance in LFN-exposed mice but not in non-exposed mice or HFN-exposed mice. Immunohistochemical analysis showed a decreased number of vestibular hair cells and increased levels of oxidative stress in LFN-exposed mice compared to those in non-exposed mice. Our results suggest that chronic exposure to LFN at moderate levels causes impaired balance involving morphological impairments of the vestibule with enhanced levels of oxidative stress. Thus, the results of this study indicate the importance of considering the risk of chronic exposure to LFN at a moderate level for imbalance.     

Velocity and attenuation of sound in the isolated fetal lung as it is expanded with air.

We measured the velocity and attenuation of audible sound in the isolated lung of the near-term fetal sheep to test the hypothesis that the acoustic properties of the lung provide a measure of the volume of gas it contains. We introduced pseudorandom noise (bandwidth 70 Hz-7 kHz) to one side of the lung and recorded the noise transmitted to the surface immediately opposite, starting with the lung containing only fetal lung liquid and making measurements after stepwise inflation with air until a leak developed. The velocity of sound in the lung fell rapidly from 187 +/- 28.2 to 87 +/- 3.7 m/s as lung density fell from 0.93 +/- 0.01 to 0.75 +/- 0.01 g/ml (lung density = lung weight/gas volume plus lung tissue volume). For technical reasons, no estimate of velocity could be made before the first air injection. Thereafter, as lung density fell to 0.35 +/- 0.01 g/ml, there was a further decline in velocity to 69.6 +/- 4.6 m/s. High-frequency sound was attenuated as lung density decreased from 1.0 to 0.5 g/ml, with little change thereafter down to a density of 0.35 +/- 0.01 g/ml. We conclude that both the velocity of audible sound through the lung and the degree to which high-frequency sound is attenuated in the lung provide information on the degree of inflation of the isolated fetal lung, particularly at high lung densities. If studies of sound transmission through the lung in the intact organism were to confirm these findings, the acoustic properties of the lung could provide a means for monitoring lung aeration during mechanical ventilation of newborn infants.     

Acoustical and neural aspects of hearing in the Australian gleaning bats,Macroderma gigas andNyctophilus gouldi

1. The maximum acoustic gain of the external ear in Macroderma gigas was found to be 25-30 dB between 5-8 kHz and in Nyctophilus gouldi it reached 15-23 dB between 7-22 kHz. Pinna gain reached a peak of 16 dB near 4.5-6 kHz in M. gigas and 12-17 dB between 7-12 kHz in N. gouldi, with average gain of 6-10 dB up to 100 kHz. Pinna gain curves resemble that of a finite conical horn, including resonance. 2. The directional properties of the external ear in both species result from sound diffraction at the pinna face, as it approximates a circular aperture. The frequency dependent movement of the acoustic axis in azimuth and elevation is attributed to the asymmetrical structure of the pinnae. 3. Evoked potentials and neuronal responses were studied in the inferior colliculus. In M. gigas, the neural audiogram has sensitivity peaks at 10-20 kHz and 35-43 kHz, with extremely low thresholds (-18 dB SPL) in the low frequency region. In N. gouldi, the neural audiogram has sensitivity peaks at 8-14 kHz (lowest threshold 5 dB SPL) and 22-45 kHz. Removal of the contralateral pinna causes a frequency dependent loss in neural threshold sensitivity of up to 10-15 dB in both species. 4. The high frequency peak in the audiogram coincides with the sonar energy band in both species, whereas the low frequency region is used for social communication. Highly sensitive low frequency hearing is discussed in relation to hunting in bats by passive listening.     

THE ULTRASONIC SONG OF THE MOTH AMYNA NATALIS (LEPIDOPTERA: NOCTUIDAE: ACONTIINAE)

ABSTRACT

In Malaysia, males of the noctuid moth Amyna natalis were observed producing a continuous ultrasonic song of high intensity (about 102 dB SPL measured at a distance of 10 cm). The frequency spectrum of the sound impulses had its peak between 60 and 80 kHz. During song production the animals were perching on plants and moving their wings up and down quickly. Simultaneously, by twisting the wings it seems likely that a male-specific “bubble” in the forewing functions as a tymbal, resulting in sound production.     

Size and scale effects as constraints in insect sound communication

For optimal transfer of power to the surrounding medium, a sound source should have a radius of 1/6 to 1/4 of the sound wavelength. Sound-waves propagate from the source as compressions and rarefactions of the fluid medium, which decay by spreading and viscous losses. Higher frequencies are more easily refracted and reflected by objects in the environment, causing degradation of signal structure. In open air or water, the sound spreads spherically and decays by the inverse square law. If the sound is restricted to two dimensions rather than three, it decays as the inverse of range, whereas waves within a rod decay largely due to viscous losses; such calls are usually rather simple pulses and rely on the initial time of arrival because of multiple pathlengths or different propagation velocities in the environment. Because of the relationship between calling success and reproductive success, singing insects are under selective pressure to optimize the range, and to maintain the specificity, of their calls. Smaller insects have less muscle power; because of their small sound sources, higher frequencies will be radiated more efficiently than lower frequencies, but in order to produce brief loud pulses from a long-duration muscle contraction they may use both a frequency multiplier mechanism and a mechanical power amplifier. Airborne insect sounds in the range from 1 to 5 kHz tend to have sustained puretone components and a specific pattern of pulses which propagate accurately. Where the song frequency is higher, the pulses tend to become briefer, with a rapid initial build-up that gives a reliable time of onset through obstructed transmission pathways. These scale effects may be related both to the sound-producing mechanism and the auditory system of the receiver. Tiny insects have the special acoustic problem of communicating with only a small amount of available power. Some, such as fruit flies, communicate at low frequencies, at close range, by generating air currents; these currents may also be used to waft specific pheromones. Other small insects, such as Hemiptera, beetles, etc., communicate using substrate vibration. This enables long-range communication, but signal structure degrades with distance from the source; vibration signals tend to be confined to certain types of linear substrate, such as vegetation.     

Azimuthal sound localization in the European starling (Sturnus vulgaris)

Summary The physical measurements reported here test whether the European starling (Sturnus vulgaris) evaluates the azimuth direction of a sound source with a peripheral auditory system composed of two acoustically coupled pressure-difference receivers (1) or of two decoupled pressure receivers (2).A directional pattern of sound intensity in the freefield was measured at the entrance of the auditory meatus using a probe microphone, and at the tympanum using laser vibrometry. The maximum differences in the soundpressure level measured with the microphone between various speaker positions and the frontal speaker position were 2.4 dB at 1 and 2 kHz, 7.3 dB at 4 kHz, 9.2 dB at 6 kHz, and 10.9 dB at 8 kHz. The directional amplitude pattern measured by laser vibrometry did not differ from that measured with the microphone. Neither did the directional pattern of travel times to the ear. Measurements of the amplitude and phase transfer function of the starling's interaural pathway using a closed sound system were in accord with the results of the free-field measurements.In conclusion, although some sound transmission via the interaural canal occurred, the present experiments support the hypothesis 2 above that the starling's peripheral auditory system is best described as consisting of two functionally decoupled pressure receivers.Abbreviations CM cochlear microphonics - ITD interaural time difference - IID interaural intensity difference - MRA minimum resolvable angle - dB SPL sound-pressure level (re 0.00002 Pa)     

大鼠听神经单纤维的活动

本文以声压级(SP)的dB值为单位,用不同频率(从音频到超声)的声刺激,对大鼠听觉一级神经元325根单一纤维的活动进行了观察。结果表明:每一纤维都有自己的最佳频率和相应的最低阈值。测得最佳频率的最低值为0.58kHz,最高值为62.6kHz; 最低阈值为6dBSPL,其相应频率为27.49kHz;最敏感的频率范围在20—50kHz。频率-阈值曲线在比最佳频率高的一侧斜度陡峭,低的一侧倾斜缓慢。频率-阈值曲线的锐度若以Q值表示,它对最佳频率分布的回归曲线由最佳频率的低频向高频方向逐渐升高,且Q10,Q20,Q30,Q40,Q50,dB的回归曲线具有相似的倾斜度。绝大多数纤维都有自发放电。给最佳频率持续音作用时,随刺激强度的增强,放电速率增加,但到阈上30dB左右皆达饱和。由各频率的最低阈值绘成的听反应阈曲线与行为测听所得的听力曲线颇为近似。     

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).     

Tympanic and extratympanic sound transmission in the leopard frog

Summary The inner ear of the leopard frog,Rana pipiens, receives sound via two separate pathways: the tympanic-columellar pathway and an extratympanic route. The relative efficiency of the two pathways was investigated. Laser interferometry measurements of tympanic vibration induced by free-field acoustic stimulation reveal a broadly tuned response with maximal vibration at 800 and 1500 Hz. Vibrational amplitude falls off rapidly above and below these frequencies so that above 2 kHz and below 300 Hz tympanic vibration is severely reduced. Electrophysiological measurements of the thresholds of single eighth cranial nerve fibers from both the amphibian and basilar papillae in response to pure tones were made in such a way that the relative efficiency of tympanic and extratympanic transmission could be assessed for each fiber. Thresholds for the two routes are very similar up to 1.0 kHz, above which tympanic transmission eventually becomes more efficient by 15–20 dB. By varying the relative phase of the two modes of stimulation, a reduction of the eighth nerve response can be achieved. When considered together, the measurements of tympanic vibration and the measurements of tympanic and extratympanic transmission thresholds suggest that under normal conditions in this species (1) below 300 Hz extratympanic sound transmission is the main source of inner ear stimulation; (2) for most of the basilar papilla frequency range (i.e., above 1.2 kHz) tympanic transmission is more important; and (3) both routes contribute to the stimulation of amphibian papilla fibers tuned between those points. Thus acoustic excitation of the an uran's inner ear depends on a complex interac tion between tympanic and extratympanic sound transmission.Abbreviations dB SPL decibels sound pressure level re: 20 N/ m² - AP amphibian papilla - BP basilar papilla - BEF best excitatory frequency     

Involvement of the mechanosensory complex structures of the cricket <Emphasis Type="Italic">Phaeophilacris bredoides</Emphasis> in triggering of motor responses to sound

Using an ethological approach, we studied the possibility of sound perception as well as probable contribution of diverse mechanosensory systems composing the mechanosensory complex to triggering of motor responses to sound stimulation in imaginal crickets Phaeophilacris bredoides lacking the tympanal organs (“deaf”). It was shown that Ph. bredoides imagoes are able to perceive sounds and respond to sound cues by a locomotor reaction in a relatively broad frequency range which becomes narrower as sound intensity decreases [0.1–6.0 kHz (111 ± 3 dB SPL), 0.1–1.5 kHz (101 ± 3 dB SPL), 0.1–1.3 kHz (91 ± 3 dB SPL), 0.1–0.6 kHz (81 ± 3 dB SPL), and 0.1 kHz (71 ± 3 dB SPL)]. Sound perception and triggering ofmotor responses appear to involve the cercal organs (CO), subgenual organs (SO) and, probably, other distant mechanosensory organs (DMO). CO are essential for triggering of locomotor responses to sound within the ranges of 1.6–6.0 kHz (111 ± 3 dB SPL), 1–1.5 kHz (101 ± 3 dB SPL), 0.9–1.3 kHz (91 ± 3 dB SPL), and 0.5–0.6 kHz (81 ± 3 dB SPL). SO and, probably, other DMO provide locomotor responses to sound within the ranges of 0.1–6.0 kHz (111 ± 3 dB SPL), 0.1–0.8 kHz (101 ± 3 dB SPL), 0.1–0.4 kHz (91 ± 3 dB SPL), and 0.1–0.4 kHz (81 ± 3 dB SPL). From this, it follows that “deaf” (nonsinging) Ph. bredoides can perceive sounds using CO, SO and, probably, other DMO, which (as in singing crickets) are likely to compose an integrated mechanosensory complex providing adequate acoustic behavior of this cricket species. Performance efficiency and sensitivity of the mechanosensory complex (specifically, of CO) rely on the thoroughness of grooming. Following self-cleaning of CO, the level of cricket motor activity in response to cue presentation returned to the baseline and sometimes even increased. Whether or not crickets of this species communicate acoustically is yet to be found out, however, we suggest that the mechanosensory complex, which triggers motor responses to a sound, is normally involved in the defensive escape response aimed at rescuing from predators.     

TEMPORARY THRESHOLD SHIFTS AFTER NOISE EXPOSURE IN THE BOTTLENOSE DOLPHIN (TURSIOPS TRUNCATUS) MEASURED USING EVOKED AUDITORY POTENTIALS

The time course of recovery from temporary threshold shift (TTS) was measured in a bottlenose dolphin, Tursiops truncatus , using an evoked-potential procedure. The envelope-following response (EFR), which is a rhythmic train of auditory brainstem responses (ABR) to sinusoidally amplitude-modulated tones, was used as an indicator of the sound reception by the animal. Variation of the intensity of the stimulus allowed us to measure the animal's hearing via EFR thresholds. During each session, following an initial measure of threshold, the trained animal voluntary positioned itself within a hoop 1 m underwater while a 160 dB re 1 μPa noise of a 4–11 kHz bandwidth was presented for 30 min. After the noise exposure, thresholds were measured again at delays of 5, 10, 15, 25, 45, and 105 min. Measurements were made at test frequencies of 8, 11.2, 16, 22.5, and 32 kHz. The maximum TTS occurred 5 min after exposure and rapidly recovered with a rate of around 1.5 dB per doubling of time. TTS occurred at test frequencies from 8 to 16 kHz, with the maximum at 16 kHz. TTS was negligible at 22.5 kHz and absent at 32 kHz.     

Physics of directional hearing in the cricket Gryllus bimaculatus

In the cricket ear, sound acts on the external surface of the tympanum and also reaches the inner surface after travelling in at least three pathways in the tracheal system. We have determined the transmission gain of the three internal sound pathways; that is, the change of amplitude and phase angle from the entrances of the tracheal system to the inner surface of the tympanum. In addition, we have measured the diffraction and time of arrival of sound at the ear and at the three entrances at various directions of sound incidence. By combining these data we have calculated how the total driving force at the tympanum depends on the direction of sound. The results are in reasonable agreement with the directionality of the tympanal vibrations as determined with laser vibrometry.At the frequency of the calling song (4.7 kHz), the direction of the sound has little effect on the amplitudes of the sounds acting on the tympanum, but large effects on their phase angles, especially of the sound waves entering the tracheal system at the contralateral side of the body. The master parameter for causing the directionality of the ear in the forward direction is the sound wave entering the contralateral thoracic spiracle. The phase of this sound component may change by 130–140° with sound direction. The transmission of sound from the contralateral inputs is dominated by a very selective high-pass filter, and large changes in amplitude and phase are seen in the transmitted sounds when the sound frequency changes from 4 to 5 kHz. The directionality is therefore very dependent on sound frequency.The transmission gains vary considerably in different individuals, and much variation was also found in the directional patterns of the ears, especially in the effects of sounds from contralateral directions. However, the directional pattern in the frontal direction is quite robust (at least 5 dB difference between the 330° and 30° directions), so these variations have only little effect on how well the individual animals can approach singing conspecifics.Abbreviations CS contralateral spiracle - CT contralateral tympanum - IS ipsilateral spiracle - IT ipsilateral tympanum - P the vectorial sum of the sounds acting on the tympanum     

Source levels and the estimated active space of bottlenose dolphin (Tursiops truncatus) whistles in the Moray Firth, Scotland

This study measured SPLs of whistles of wild bottlenose dolphins (Tursiops truncatus) in the Moray Firth, Scotland, and estimated their active space, i.e. the distance at which another dolphin can perceive the whistle of a conspecific. Whistling dolphins were localized with a dispersed hydrophone array by comparing differences in the times of arrival of a whistle at different hydrophones. The mean source level for whistles was 158 +/- 0.6 dB re. 1 microPa. The maximum was 169 dB re. 1 microPa. The active space of these whistles was calculated taking into account transmission loss, ambient noise, the critical ratios and the auditory sensitivity of this species. The estimated radius of the active space of unmodulated whistles between 3.5 kHz and 10 kHz produced at maximum source level ranged from 20 km to 25 km in a habitat of 10 m depth and at sea state 0. At sea state 4 it ranged from 14 km to 22 km. For whistles of 12 kHz it dropped to 1.5-4 km. The results suggest that whistles can be used to maintain group cohesion over large distances but also that dolphins that researchers consider to belong to separate groups might be in acoustic contact.