Temporary threshold shift in bottlenose dolphins (Tursiops truncatus) exposed to mid-frequency tones

A behavioral response paradigm was used to measure hearing thresholds in bottlenose dolphins before and after exposure to 3 kHz tones with sound exposure levels (SELs) from 100 to 203 dB re 1 microPa2 s. Experiments were conducted in a relatively quiet pool with ambient noise levels below 55 dB re 1 microPa2/Hz at frequencies above 1 kHz. Experiments 1 and 2 featured 1-s exposures with hearing tested at 4.5 and 3 kHz, respectively. Experiment 3 featured 2-, 4-, and 8-s exposures with hearing tested at 4.5 kHz. For experiment 2, there were no significant differences between control and exposure sessions. For experiments 1 and 3, exposures with SEL=197 dB re 1 microPa2 s and SEL > or = 195 dB re 1 microPa2 s, respectively, resulted in significantly higher TTS4 than control sessions. For experiment 3 at SEL= 195 dB re 1 microPa2 s, the mean TTS4 was 2.8 dB. These data are consistent with prior studies of TTS in dolphins exposed to pure tones and octave band noise and suggest that a SEL of 195 dB re 1 microPa2 s is a reasonable threshold for the onset of TTS in dolphins and white whales exposed to midfrequency tones.     

Assessing temporary threshold shift in a bottlenose dolphin (Tursiops truncatus) using multiple simultaneous auditory evoked potentials

Hearing sensitivity was measured in a bottlenose dolphin before and after exposure to an intense 20-kHz fatiguing tone in three different experiments. In each experiment, hearing was characterized using both the auditory steady-state response (ASSR) and behavioral methods. In experiments 1 and 2, ASSR stimuli consisted of seven frequency-modulated tones, each with a unique carrier and modulation frequency. The tones were simultaneously presented to the subject and the ASSR at each modulation rate measured to determine the effects of the sound exposure at the corresponding carrier frequency. In experiment 3 behavioral thresholds and ASSR input-output functions were measured at a single frequency before and after three exposures. Hearing loss was frequency-dependent, with the largest temporary threshold shifts occurring (in order) at 30, 40, and 20 kHz. ASSR threshold shifts reached 40-45 dB and were always larger than behavioral shifts (19-33 dB). The ASSR input-output functions were represented as the sum of two processes: a low threshold, saturating process and a higher threshold, linear process, that react and recover to fatigue at different rates. The loss of the near-threshold saturating process after exposure may explain the discrepancies between the ASSR and behavioral threshold shifts.     

The biosonar field around an Atlantic bottlenose dolphin (Tursiops truncatus)

The use of remote autonomous passive acoustic recorders (PAR) to determine the distribution of dolphins at a given locations has become very popular. Some investigators are using echolocation clicks to gather information on the presence of dolphins and to identify species. However, in all of these cases, the PAR probably recorded mainly off-axis clicks, even some from behind the animals. Yet there is a very poor understanding of the beam pattern and the click waveform and spectrum from different azimuths around the animal's body. The beam pattern completely around an echo locating dolphin was measured at 16 different but equally spaced angles in the horizontal plane using an 8-hydrophone array in sequence. Eight channels of data were digitized simultaneously at a sampling rate of 500 kHz. The resulting beam patterns in both planes showed a continuous drop off in sound pressure with azimuth around the animal and reached levels below -50 dB relative to the signal recorded on the beam axis. The signals began to break up into two components at angles greater than ± 45° in the horizontal plane. The center frequency dropped off from its maximum at 0° in a non-uniform matter.     

Simultaneously measured behavioral and electrophysiological hearing thresholds in a bottlenose dolphin (Tursiops truncatus)

Dolphin auditory thresholds obtained via evoked potential audiometry may deviate from behavioral estimates by 20 dB or more. Differences in the sound source, stimulus presentation method, wave form, and duration may partially explain these discrepancies. To determine the agreement between behavioral and auditory evoked potential (AEP) threshold estimates when these parameters are held constant, behavioral and AEP hearing tests were simultaneously conducted in a bottlenose dolphin. Measurements were made in-air, using sinusoidal amplitude-modulated tones continuously projected via a transducer coupled to the pan region of the dolphin's lower jaw. Tone trials were presented using the method of constant stimuli. Behavioral thresholds were estimated using a 50% correct detection. AEP thresholds were based on the envelope following response and 50% correct detection. Differences between AEP and behavioral thresholds were within +/-5 dB, except at 10 kHz (12 dB), 20 kHz (8 dB), 30 kHz (7 dB), and 150 kHz (24 dB). In general, behavioral thresholds were slightly lower, though this trend was not significant. The results demonstrate that when the test environment, sound source, stimulus wave form, duration, presentation method, and analysis are consistent, the magnitude of the differences between AEP and behavioral thresholds is substantially reduced.     

Temporary threshold shifts and recovery following noise exposure in the Atlantic bottlenosed dolphin (Tursiops truncatus)

Behaviorally determined hearing thresholds for a 7.5-kHz tone for an Atlantic bottlenosed dolphin (Tursiops truncatus) were obtained following exposure to fatiguing low-frequency octave band noise. The fatiguing stimulus ranged from 4 to 11 kHz and was gradually increased in intensity to 179 dB re 1 microPa and in duration to 55 min. Exposures occurred no more frequently than once per week. Measured temporary threshold shifts averaged 11 dB. Threshold determination took at least 20 min. Recovery was examined 360, 180, 90, and 45 min following exposure and was essentially complete within 45 min.     

Whole-lung resonance in a bottlenose dolphin (Tursiops truncatus) and white whale (Delphinapterus leucas)

An acoustic backscatter technique was used to estimate in vivo whole-lung resonant frequencies in a bottlenose dolphin (Tursiops truncatus) and white whale (Delphinapterus leucas). Subjects were trained to submerge and position themselves near an underwater sound projector and a receiving hydrophone. Acoustic pressure measurements were made near the thorax while the subject was insonified with pure tones at frequencies from 16 to 100 Hz. Whole-lung resonant frequencies were estimated by comparing pressures measured near the subject's thorax to those measured from the same location without the subject present. Experimentally measured resonant frequencies for the white whale and dolphin lungs were 30 and 36 Hz, respectively. These values were significantly higher than those predicted using a free-spherical air bubble model. Experimentally measured damping ratios and quality factors at resonance were 0.20 and 2.5, respectively, for the white whale, and 0.16 and 3.1, respectively, for the dolphin.     

Target detection in reverberation by an echolocating Atlantic bottlenose dolphin (Tursiops truncatus)

An experiment was conducted to determine the ability of an echolocating Atlantic bottlenose dolphin (Tursiops truncatus) to detect targets in the presence of reverberation. Reverberation was induced by a clutter screen consisting of forty-eight 5.1-cm-diam cork spheres spaced 15.2 cm apart and arranged in a rectangular array, located behind the targets. Hollow aluminum cylinders having the same outer diameter and wall thickness and three different lengths were used as targets. The dolphin was trained to station in a hoop, 6 m from the targets, and to echolocate the target upon the reception of an audio cue. Only one of the targets would be presented in a target-present trial. Data were collected on the animal's detection performance as a function of the separation distance between the clutter screen and the targets. The animal's performance for the smallest cylinder varied from 91% to 55% correct as the separation distance decreased from 10.2 to 0 cm. The target strength of the clutter screen and the cylinders were measured both in terms of the energies and the maximum peak-to-peak amplitudes of the incident and reflected echoes. These measurements indicated that when the target-clutter screen separation distance was 0 cm, the dolphin's 50% correct detection occurred at an energy echo-to-reverberation ratio of approximately 0.25 dB and at peak-to-peak echo-to-reverberation ratio of 2.6 dB. The results also indicated that the dolphin's performance varied almost linearly with the echo-to-reverberation ratio. The animal's pulse emissions were monitored by a microprocessor system and the results are presented in terms of the average number of clicks and the average response latency, as a function of the separation distance.     

Subjective loudness level measurements and equal loudness contours in a bottlenose dolphin (Tursiops truncatus)

Loudness level measurements in human listeners are straightforward; however, it is difficult to convey the concepts of loudness matching or loudness comparison to (non-human) animals. For this reason, prior studies have relied upon objective measurements, such as response latency, to estimate equal loudness contours in animals. In this study, a bottlenose dolphin was trained to perform a loudness comparison test, where the listener indicates which of two sequential tones is louder. To enable reward of the dolphin, most trials featured tones with identical or similar frequencies, but relatively large sound pressure level differences, so that the loudness relationship was known. A relatively small percentage of trials were "probe" trials, with tone pairs whose loudness relationship was not known. Responses to the probe trials were used to construct psychometric functions describing the loudness relationship between a tone at a particular frequency and sound pressure level and that of a reference tone at 10 kHz with a sound pressure level of 90, 105, or 115 dB re 1 μPa. The loudness relationships were then used to construct equal loudness contours and auditory weighting functions that can be used to predict the frequency-dependent effects of noise on odontocetes.     

Bottlenose dolphin (Tursiops truncatus) steady-state evoked responses to multiple simultaneous sinusoidal amplitude modulated tones

Auditory steady-state evoked potentials were measured in a bottlenose dolphin (Tursiops truncatus) in response to single and multiple sinusoidal amplitude modulated (SAM) tones. Tests were conducted in air using a "jawphone" sound projector. Evoked potentials were recorded noninvasively using surface electrodes embedded in suction cups. Sound stimuli consisted of SAM tones with 1, 2, 3, or 4 carrier frequencies (10, 20, 30, 40 kHz), each with a unique modulation frequency. Stimulus sound pressure levels were varied in 5-dB steps from approximately 120 to 60-75 dB re 1 microPa, depending on frequency. Evoked potentials followed the temporal envelope of each stimulus, resulting in spectral components at each unique modulation frequency. Spectral analysis was used to evaluate the response amplitude for each carrier as a function of stimulus level. There were no significant differences between thresholds obtained with single and multiple stimuli at 10, 30, and 40 kHz. At 20 kHz, thresholds obtained with three components were higher than those obtained with four components, possibly revealing interactions between stimuli with less than one octave frequency separation. The use of multiple SAM stimuli may offer substantial advantages for studies of marine mammal hearing, where testing time and access to subjects are typically limited.     

Underwater sound pressure variation and bottlenose dolphin (Tursiops truncatus) hearing thresholds in a small pool

Studies of underwater hearing are often hampered by the behavior of sound waves in small experimental tanks. At lower frequencies, tank dimensions are often not sufficient for free field conditions, resulting in large spatial variations of sound pressure. These effects may be mitigated somewhat by increasing the frequency bandwidth of the sound stimulus, so effects of multipath interference average out over many frequencies. In this study, acoustic fields and bottlenose dolphin (Tursiops truncatus) hearing thresholds were compared for pure tone and frequency modulated signals. Experiments were conducted in a vinyl-walled, seawater-filled pool approximately 3.7 x 6 x 1.5 m. Acoustic signals were pure tone and linear and sinusoidal frequency modulated tones with bandwidths/modulation depths of 1%, 2%, 5%, 10%, and 20%. Thirteen center frequencies were tested between 1 and 100 kHz. Acoustic fields were measured (without the dolphin present) at three water depths over a 60 x 65 cm grid with a 5-cm spacing. Hearing thresholds were measured using a behavioral response paradigm and up/down staircase technique. The use of FM signals significantly improved the sound field without substantially affecting the measured hearing thresholds.     

Bio-inspired wideband sonar signals based on observations of the bottlenose dolphin (Tursiops truncatus)

This paper uses advanced time-frequency signal analysis techniques to generate new models for bio-inspired sonar signals. The inspiration comes from the analysis of bottlenose dolphin clicks. These pulses are very short duration, between 50 and 80 micros, but for certain examples we can delineate a double down-chirp structure using fractional Fourier methods. The majority of clicks have energy distributed between two main frequency bands with the higher frequencies delayed in time by 5-20 micros. Signal syntheses using a multiple chirp model based on these observations are able to reproduce much of the spectral variation seen in earlier studies on natural dolphin echolocation pulses. Six synthetic signals are generated and used to drive the dolphin based sonar (DBS) developed through the Biosonar Program office at the SPAWAR Systems Center, San Diego, CA. Analyses of the detailed echo structure for these pulses ensonifying two solid copper spherical targets indicate differences in discriminatory potential between the signals. It is suggested that target discrimination could be improved through the transmission of a signal packet in which the chirp structure is varied between pulses. Evidence that dolphins may use such a strategy themselves comes from observations of variations in the transmissions of dolphins carrying out target detection and identification tasks.     

Temporary shift in masked hearing thresholds of bottlenose dolphins, Tursiops truncatus, and white whales, Delphinapterus leucas, after exposure to intense tones

A behavioral response paradigm was used to measure masked underwater hearing thresholds in five bottlenose dolphins and two white whales before and immediately after exposure to intense 1-s tones at 0.4, 3, 10, 20, and 75 kHz. The resulting levels of fatiguing stimuli necessary to induce 6 dB or larger masked temporary threshold shifts (MTTSs) were generally between 192 and 201 dB re: 1 microPa. The exceptions occurred at 75 kHz, where one dolphin exhibited an MTTS after exposure at 182 dB re: 1 microPa and the other dolphin did not show any shift after exposure to maximum levels of 193 dB re: 1 microPa, and at 0.4 kHz, where no subjects exhibited shifts at levels up to 193 dB re: 1 microPa. The shifts occurred most often at frequencies above the fatiguing stimulus. Dolphins began to exhibit altered behavior at levels of 178-193 dB re: 1 microPa and above; white whales displayed altered behavior at 180-196 dB re: 1 microPa and above. At the conclusion of the study all thresholds were at baseline values. These data confirm that cetaceans are susceptible to temporary threshold shifts (TTS) and that small levels of TTS may be fully recovered.     

Directional properties of bottlenose dolphin (Tursiops truncatus) clicks, burst-pulse, and whistle sounds

The directional properties of bottlenose dolphin clicks, burst-pulse, and whistle signals were measured using a five element array, at horizontal angles of 0°, 45°, 90°, 135°, and 180° relative to a dolphin stationed on an underwater biteplate. Clicks and burst-pulse signals were highly directional with directivity indices of ~11 dB for both signal types. Higher frequencies and higher amplitudes dominated the forward, on-axis sound field. A similar result was found with whistles, where higher frequency harmonics had greater directivity indices than lower frequency harmonics. The results suggest the directional properties of these signals not only provide enhanced information to the sound producer (as in echolocation) but can provide valuable information to conspecific listeners during group coordination and socialization.     

Comodulation masking release in bottlenose dolphins (Tursiops truncatus)

The acoustic environment of the bottlenose dolphin often consists of noise where energy across frequency regions is coherently modulated in time (e.g., ambient noise from snapping shrimp). However, most masking studies with dolphins have employed random Gaussian noise for estimating patterns of masked thresholds. The current study demonstrates a pattern of masking where temporally fluctuating comodulated noise produces lower masked thresholds (up to a 17 dB difference) compared to Gaussian noise of the same spectral density level. Noise possessing wide bandwidths, low temporal modulation rates, and across-frequency temporal envelope coherency resulted in lower masked thresholds, a phenomenon known as comodulation masking release. The results are consistent with a model where dolphins compare temporal envelope information across auditory filters to aid in signal detection. Furthermore, results suggest conventional models of masking derived from experiments using random Gaussian noise may not generalize well to environmental noise that dolphins actually encounter.     

Estimating bottlenose dolphin (Tursiops truncatus) hearing thresholds from single and multiple simultaneous auditory evoked potentials

Hearing thresholds were estimated in four bottlenose dolphins by measuring auditory evoked responses to single and multiple sinusoidal amplitude modulated tones. Subjects consisted of two males and two females with ages from 4 to 22 years. Testing was conducted in air using a "jawphone" transducer to couple sound into each subject's lower right jaw. Carrier frequencies ranged from 10 to 160 kHz in one-half octave steps. Amplitude modulated stimuli were presented individually and as the sum of four, five, and nine simultaneous tones with unique carrier and modulation frequencies. Evoked potentials were noninvasively recorded using surface electrodes embedded in silicon suction cups. The presence or absence of an evoked response at each modulation frequency was assessed by calculating the magnitude-squared coherence from the frequency spectra of the recorded sweeps. All subjects exhibited traditional "U-shaped" audiograms with upper cutoff frequencies above 113 kHz. The time required for threshold estimates ranged from 23 to 37 min for single stimuli to 5-9 min for nine simultaneous stimuli. Agreement between thresholds estimated from single stimuli and multiple, simultaneous stimuli was generally good, indicating that multiple stimuli may be used for quick hearing assessment when time is limited.     

Synthesis and modification of the whistles of the bottlenose dolphin, Tursiops truncatus

A signal-processing algorithm was developed to analyze harmonic frequency-modulated sounds, to modify the parameters of the analyzed signal, and to synthesize a new analytically specified signal that resembles the original signal in specified features. This algorithm was used with dolphin whistles, a frequency-modulated harmonic signal that has typically been described in terms of its contour, or pattern of modulation of the fundamental frequency. In order to test whether other features may also be salient to dolphins, the whistle analysis calculates the energies at the harmonics as well as the fundamental frequency of the whistle. The modification part of the algorithm can set all of these energies to a constant, can shift the whistle frequency, and can expand or compress the time base or the frequency of the whistle. The synthesis part of the algorithm then synthesizes a waveform based upon the energies and frequencies of the fundamental and first two harmonics. These synthetic whistles will be useful for evaluating what acoustic features dolphins use in discriminating different whistles.     

Frequency and level dependent masking of the multiple auditory steady-state response in the bottlenose dolphin (Tursiops truncatus)

The potential for interactions between steady-state evoked responses to simultaneous auditory stimuli was investigated in two bottlenose dolphins (Tursiops truncatus). Three experiments were conducted using either a probe stimulus (probe condition) or a probe in the presence of a masker (probe-plus-masker condition). In the first experiment, the probe and masker were sinusoidal amplitude-modulated (SAM) tones. Probe and masker frequencies and masker level were manipulated to provide variable masking conditions. Probe frequencies were 31.7, 63.5, 100.8, and 127.0 kHz. The second experiment was identical to the first except only the 63.5 kHz probe was used and maskers were pure tones. For the third experiment, thresholds were measured for the probe and probe-plus-masker conditions using two techniques, one based on the lowest detectable response and the other based on a regression analysis. Results demonstrated localized masking effects where lower frequency maskers suppressed higher frequency probes and higher amplitude maskers produced a greater masking effect. The pattern of pure tone masking was nearly identical to SAM tone masking. The two threshold estimates were similar in low masking conditions, but in high masking conditions the lowest detectable response tended to overestimate thresholds while the regression-based analysis tended to underestimate thresholds.     

Temporary threshold shift in human subject exposed to noise

Using an audiometer,the effect of the noise level upon temporarythreshold shift(TTS)for five trained normal subjects(left ear only)was studied.The measurements were carried out after 6 min exposure(in third octave band)for different sound pressure levels ranging between 75-105 dB at three test fre-quencies 2,3,and 4 kHz.The results indicated that at exposure to noise of soundpressure level(SPL)above 85 dB,TTS increases linearly with ths SPL for all thetest frequencies.The work had extended to study the recovery curves for the sameears.The results indicated that the reduction in TTS on doubling the recoverytimes,for the two sound pressure levels 95 dB and 105 dB,occurs at a rate of near-ly 3 dB.The comparison of the recovery curve at 3 kHz with that calculated usingWard's general equation for recovery was made.Finally,to study the values ofTTS produced by exposure to certain noise at different test frequencies,distribu-tion curves for two recovery times were plotted representing TTS values,for anexposure     

Auditory filter shapes for the bottlenose dolphin (Tursiops truncatus) and the white whale (Delphinapterus leucas) derived with notched noise

Auditory filter shapes were estimated in two bottlenose dolphins (Tursiops truncatus) and one white whale (Delphinapterus leucas) using a behavioral response paradigm and notched noise. Masked thresholds were measured at 20 and 30 kHz. Masking noise was centered at the test tone and had a bandwidth of 1.5 times the tone frequency. Half-notch width to center frequency ratios were 0, 0.125, 0.25, 0.375, and 0.5. Noise spectral density levels were 90 and 105 dB re: 1 microPa2/Hz. Filter shapes were approximated using a roex(p,r) function; the parameters p and r were found by fitting the integral of the roex(p,r) function to the measured threshold data. Mean equivalent rectangular bandwidths (ERBs) calculated from the filter shapes were 11.8 and 17.1% of the center frequency at 20 and 30 kHz, respectively, for the dolphins and 9.1 and 15.3% of the center frequency at 20 and 30 kHz, respectively, for the white whale. Filter shapes were broader at 30 kHz and 105 dB re: 1 microPa2/Hz masking noise. The results are in general agreement with previous estimates of ERBs in Tursiops obtained with a behavioral response paradigm.     

Source parameters of echolocation clicks from wild bottlenose dolphins (Tursiops aduncus and Tursiops truncatus)

The Indian Ocean and Atlantic bottlenose dolphins (Tursiops aduncus and Tursiops truncatus) are among the best studied echolocating toothed whales. However, almost all echolocation studies on bottlenose dolphins have been made with captive animals, and the echolocation signals of free-ranging animals have not been quantified. Here, biosonar source parameters from wild T. aduncus and T. truncatus were measured with linear three- and four-hydrophone arrays in four geographic locations. The two species had similar source parameters, with source levels of 177-228 dB re 1 μPa peak to peak, click durations of 8-72 μs, centroid frequencies of 33-109 kHz and rms bandwidths between 23 and 54 kHz. T. aduncus clicks had a higher frequency emphasis than T. truncatus. The transmission directionality index was up to 3 dB higher for T. aduncus (29 dB) as compared to T. truncatus (26 dB). The high directionality of T. aduncus does not appear to be only a physical consequence of a higher frequency emphasis in clicks, but may also be caused by differences in the internal properties of the sound production system.