Temporal integration of tone signals by a killer whale (Orcinus orca).

A psychophysical procedure was used to measure pure-tone detection thresholds for a killer whale (Orcinus orca) as a function of both signal frequency and signal duration. Frequencies ranged between 1 and 100 kHz and signal durations ranged from 50 µs to 2 s, depending on the frequency. Detection thresholds decreased with an increase in signal duration up to a critical duration, which represents the auditory integration time. Integration times ranged from 4 ms at 100 kHz and increased up to 241 ms at 1 kHz. The killer whale data are similar to other odontocete species that have participated in similar experiments. The results have implications for noise impact predictions for signals with durations less than the auditory integration time.

Mechanisms of auditory masking in marine mammals.

Anthropogenic noise is an increasing threat to marine mammals that rely on sound for communication, navigation, detecting prey and predators, and finding mates. Auditory masking is one consequence of anthropogenic noise, the study of which is approached from multiple disciplines including field investigations of animal behavior, noise characterization from in-situ recordings, computational modeling of communication space, and hearing experiments conducted in the laboratory. This paper focuses on laboratory hearing experiments applying psychophysical methods, with an emphasis on the mechanisms that govern auditory masking. Topics include tone detection in simple, complex, and natural noise; mechanisms for comodulation masking release and other forms of release from masking; the role of temporal resolution in auditory masking; and energetic vs informational masking.

Spatial acuity of the bottlenose dolphin (Tursiops truncatus) biosonar system with a bat and human comparison.

Horizontal angular resolution was measured in two bottlenose dolphins using a two-alternative forced-choice, biosonar target discrimination paradigm. The task required a stationary dolphin positioned in a hoop to discriminate two physical targets at a range of 4 m. The angle separating the targets was manipulated to estimate an angular discrimination threshold of 1.5°. In a second experiment, a similar two-target biosonar discrimination task was conducted with one free-swimming dolphin, to test whether its emission beam was a critical factor in discriminating the targets. The spatial separation between two targets was manipulated to measure a discrimination threshold of 6.7 cm. There was a relationship between differences in acoustic signals received at each target and the dolphin's performance. The results of the angular resolution experiment were in good agreement with measures of the minimum audible angle of both dolphins and humans and remarkably similar to measures of angular difference discrimination in echolocating dolphins, bats, and humans. The results suggest that horizontal auditory spatial acuity may be a common feature of the mammalian auditory system rather than a specialized feature exclusive to echolocating auditory predators.

Auditory masking in killer whales (Orcinus orca): Critical ratios for tonal signals in Gaussian noise.

Masked detection thresholds were measured for two killer whales (Orcinus orca) using a psychoacoustic, adaptive-staircase procedure. Noise bands were 1-octave wide continuous Gaussian noise. Tonal signals extended between 500 Hz and 80 kHz. Resulting critical ratios increased with the signal frequency from 15 dB at 500 Hz up to 32 dB at 80 kHz. Critical ratios for killer whales were similar to those of other odontocetes despite considerable differences in size, hearing morphology, and hearing sensitivity between species.

Spectral cues and temporal integration during cylinder echo discrimination by bottlenose dolphins (Tursiops truncatus).

Three bottlenose dolphins (Tursiops truncatus) participated in simulated cylinder wall thickness discrimination tasks utilizing electronic "phantom" echoes. The first experiment resulted in psychometric functions (percent correct vs wall thickness difference) similar to those produced by a dolphin performing the task with physical cylinders. In the second experiment, a wide range of cylinder echoes was simulated, with the time separation between echo highlights covering a range from <30 to >300 µs. Dolphin performance and a model of the dolphin auditory periphery suggest that the dolphins used high-frequency, spectral-profiles of the echoes for discrimination and that the utility of spectral cues degraded when the time separation between echo highlights approached and exceeded the dolphin's temporal integration time of ∼264 µs.

Dolphin echo-delay resolution measured with a jittered-echo paradigm.

Biosonar echo delay resolution was investigated in four bottlenose dolphins (Tursiops truncatus) using a "jittered" echo paradigm, where dolphins discriminated between electronic echoes with fixed delay and those whose delay alternated (jittered) on successive presentations. The dolphins performed an echo-change detection task and produced a conditioned acoustic response when detecting a change from non-jittering echoes to jittering echoes. Jitter delay values ranged from 0 to 20 µs. A passive listening task was also conducted, where dolphins listened to simulated echoes and produced a conditioned acoustic response when signals changed from non-jittering to jittering. Results of the biosonar task showed a mean jitter delay threshold of 1.3 µs and secondary peaks in error functions suggestive of the click autocorrelation function. When echoes were jittered in polarity and delay, error functions shifted by approximately 5 µs and all dolphins discriminated echoes that jittered only in polarity. Results were qualitatively similar to those from big brown bats (Eptesicus fuscus) and indicate that the dolphin biosonar range estimator is sensitive to echo phase information. Results of the passive listening task suggested that the dolphins could not passively detect changes in timing and polarity of simulated echoes.

Directional hearing sensitivity for 2-30 kHz sounds in the bottlenose dolphin (Tursiops truncatus).

Bottlenose dolphins (Tursiops truncatus) depend on sounds at frequencies lower than 30 kHz for social communication, but little information on the directional dependence of hearing thresholds for these frequencies exists. This study measured underwater behavioral hearing thresholds for 2, 10, 20, and 30 kHz sounds projected from eight different positions around dolphins in both the horizontal and vertical planes. The results showed that the sound source direction relative to the dolphin affected hearing threshold, and that directional characteristics of the receiving beam pattern were frequency dependent. Hearing thresholds obtained from two adult dolphins demonstrated a positive relationship between directivity of hearing and stimulus frequency, with asymmetric receiving beam patterns in both the horizontal and vertical planes. Projecting sound from directly behind the dolphin resulted in frequency-dependent increases in hearing threshold up to 18.5 dB compared to when sound was projected in front. When the projector was situated above the dolphin thresholds were approximately 8 dB higher as compared to below. This study demonstrates that directional hearing exists for lower frequencies than previously expected.

The effect of jamming stimuli on the echolocation behavior of the bottlenose dolphin, Tursiops truncatus.

Echolocating bats and odontocetes face the potential challenge of acoustic interference from neighbors, or sonar jamming. To counter this, many bat species have adapted jamming avoidance strategies to improve signal detection, but any such avoidance strategies in dolphins is unknown. This study provides an investigation into whether dolphins modify echolocation behavior during jamming scenarios. Recorded echolocation clicks were projected at different click repetition rates and at different aspect angles relative to two dolphins' heads while each dolphin was performing a target detection task. Changes in the timing, amplitude, and frequency of structure of the dolphin's emitted signals were compared to determine if and how dolphins modify echolocation when faced with potentially interfering conspecific echolocation signals. The results indicate that both dolphins demonstrated different responses when faced with jamming scenarios, which may reflect optimal strategies according to individual auditory perception abilities.

Effects of vibratory pile driver noise on echolocation and vigilance in bottlenose dolphins (Tursiops truncatus).

Vibratory pile drivers, used for marine construction, can produce sustained, high sound pressure levels (SPLs) in areas that overlap with dolphin habitats. Dolphins rely on echolocation for navigation, detecting predators and prey, and to coordinate group behavior. This study examined the effects of vibratory pile driver noise on dolphin sustained target detection capabilities through echolocation. Five dolphins were required to scan their enclosure and indicate the occurrences of phantom echoes during five different source levels of vibratory pile driver playback sound (no-playback control, 110, 120, 130, and 140 dB re 1 µPa). Three of the dolphins demonstrated a significant decrease in target detection performance at 140 dB playback level that was associated with an almost complete secession of echolocation activity. The performance of two dolphins was not affected. All dolphins rapidly returned to baseline levels of target detection performance by their second replication. However, an increased number of clicks was produced at the highest playback SPL. The data suggest that the decrease in vigilant behavior was due to the vibratory pile driver noise distracting the dolphins and decreasing their motivation to perform the task.

Composite critical ratio functions for odontocete cetaceans.

Critical ratios (CRs) are useful for estimating detection thresholds of tonal signals when the spectral density of noise is known. In cetaceans, CRs have only been measured for a few animals representing four odontocete species. These data are sparse, particularly for lower frequencies where anthropogenic noise is concentrated. There is currently no systematic method for implementing CR predictions (e.g., a composite frequency-dependent CR function). The current study measures CRs for two bottlenose dolphins (Tursiops truncatus) and estimates composite CR functions. The composite models can aid in predicting and extrapolating auditory masking for a broad range of frequencies.

Killer whale (Orcinus orca) behavioral audiograms.

Killer whales (Orcinus orca) are one of the most cosmopolitan marine mammal species with potential widespread exposure to anthropogenic noise impacts. Previous audiometric data on this species were from two adult females [Szymanski, Bain, Kiehl, Pennington, Wong, and Henry (1999). J. Acoust. Soc. Am. 108, 1322-1326] and one sub-adult male [Hall and Johnson (1972). J. Acoust. Soc. Am. 51, 515-517] with apparent high-frequency hearing loss. All three killer whales had best sensitivity between 15 and 20 kHz, with thresholds lower than any odontocete tested to date, suggesting this species might be particularly sensitive to acoustic disturbance. The current study reports the behavioral audiograms of eight killer whales at two different facilities. Hearing sensitivity was measured from 100 Hz to 160 kHz in killer whales ranging in age from 12 to 52 year. Previously measured low thresholds at 20 kHz were not replicated in any individual. Hearing in the killer whales was generally similar to other delphinids, with lowest threshold (49 dB re 1 µPa) at approximately 34 kHz, good hearing (i.e., within 20 dB of best sensitivity) from 5 to 81 kHz, and low- and high-frequency hearing cutoffs (>100 dB re µPa) of 600 Hz and 114 kHz, respectively.

Energetic and informational masking of complex sounds by a bottlenose dolphin (Tursiops truncatus).

With few exceptions, laboratory studies of auditory masking in marine mammals have been limited to examining detection thresholds for simple tonal signals embedded in broadband noise. However, detection of a sound has little adaptive advantage without the knowledge of what produced the sound (recognition) and where the sound originated (localization). In the current study, a bottlenose dolphin's masked detection thresholds (energetic masking) and masked recognition thresholds (informational masking) were estimated for a variety of complex signals including dolphin vocalizations, frequency modulated signals, and a 10 kHz pure tone. Broadband noise types included recordings of natural sounds and computer generated sounds. Detection thresholds were estimated using a standard go, no-go adaptive staircase procedure. The same dolphin learned to associate whistle-like FM sounds with specific arbitrary objects using a three alternative, matching-to-sample (MTS) procedure. The dolphin's performance in the MTS task was then tested in the presence of the same masking noise types used in the detection task. Recognition thresholds were, on average, about 4 dB higher than detection thresholds for similar signal-noise conditions. The 4 dB difference is likely due to additional cognitive demands of recognition, including attention and pattern recognition.

Nearfield and farfield measurements of dolphin echolocation beam patterns: No evidence of focusing.

The potential for bottlenose dolphins to actively focus their biosonar transmissions was examined by measuring emitted clicks in four dolphins using horizontal, planar hydrophone arrays. Two hydrophone configurations were used: a rectangular array with hydrophones 0.2 to 2 m from the dolphins and a polar array with hydrophones 0.5 to 5 m from the dolphins. The biosonar task was a target change detection utilizing physical targets at ranges from 1.3 to 6.3 m with all subjects and "phantom" targets at simulated ranges from 2.5 to 20 m with two subjects. To provide a basis for evaluating the experimental data, sound fields radiated from flat and focused circular pistons were mathematically simulated using transient excitation functions similar to dolphin clicks. The array measurements showed no evidence that the dolphins adaptively focused their click emissions; axial amplitudes and iso-amplitude contours matched the pattern of the simulation results for flat transducers and showed a single region of maximum amplitude, beyond which spherical spreading loss was approximated.

Auditory sensitivity in aquatic animals.

A critical concern with respect to marine animal acoustics is the issue of hearing "sensitivity," as it is widely used as a criterion for the onset of noise-induced effects. Important aspects of research on sensitivity to sound by marine animals include: uncertainties regarding how well these species detect and respond to different sounds; the masking effects of man-made sounds on the detection of biologically important sounds; the question how internal state, motivation, context, and previous experience affect their behavioral responses; and the long-term and cumulative effects of sound exposure. If we are to better understand the sensitivity of marine animals to sound we must concentrate research on these questions. In order to assess population level and ecological community impacts new approaches can possibly be adopted from other disciplines and applied to marine fauna.

Dynamics of biosonar signals in free-swimming and stationary dolphins: The role of source levels on the characteristics of the signals.

The biosonar signals of two free-swimming Atlantic bottlenose dolphins performing a complex sonar search for a bottom target in San Diego Bay were compared with the biosonar signals of a dolphin performing a target discrimination task in a net pen in the same bay. A bite-plate device carried by the free-swimming dolphins supported a hydrophone that extended directly in front of the dolphin. A biosonar measuring tool attached to the bite plate measured the outgoing biosonar signals while the dolphins conducted sonar searches. Each of the free-swimming dolphins used different biosonar search strategy in solving the problem and the dolphins' biosonar signals reflect the difference in strategy. The dolphin in the pen stationed in a hoop while echolocating on a target 6 m away and reported if the indentation on a spherical target was directed toward it. The signals were parameterized by determining the peak-to-peak source levels, source energy flux density, peak frequency, center frequency, root-mean-square (rms) bandwidth, rms duration, and the Q of the signals. Some parameters were similar for the free-swimming and stationary dolphins while some were significantly different, suggesting biosonar signals used by free-swimming animals may be different than signals used by dolphins in a pen.

Estimating colony sizes of emerging bats using acoustic recordings.

The decline of bats demands more widespread monitoring of populations for conservation and management. Current censusing methods are either prone to bias or require costly equipment. Here, we report a new method using passive acoustics to determine bat count census from overall acoustic amplitude of the emerging bat stream. We recorded the video and audio of an emerging colony of Mexican free-tailed bats from two cave locations across multiple nights. Instantaneous bat counts were calculated from the video frames, and the bat stream's acoustic amplitude corresponding to each video frame was determined using three different methods for calculating acoustic intensity. We found a significant link between all three acoustic parameters and bat count, with the highest R (2) of 0.742 linking RMS pressure and bat count. Additionally, the relationship between acoustics and population size at one cave location could accurately predict the population size at another cave location. The data were gathered with low-cost, easy-to-operate equipment, and the data analysis can be easily accomplished using automated scripts or with open-source acoustic software. These results are a potential first step towards creating an acoustic model to estimate bat population at large cave colonies worldwide.

Recognition of Frequency Modulated Whistle-Like Sounds by a Bottlenose Dolphin (Tursiops truncatus) and Humans with Transformations in Amplitude, Duration and Frequency.

Bottlenose dolphins (Tursiops truncatus) use the frequency contour of whistles produced by conspecifics for individual recognition. Here we tested a bottlenose dolphin's (Tursiops truncatus) ability to recognize frequency modulated whistle-like sounds using a three alternative matching-to-sample paradigm. The dolphin was first trained to select a specific object (object A) in response to a specific sound (sound A) for a total of three object-sound associations. The sounds were then transformed by amplitude, duration, or frequency transposition while still preserving the frequency contour of each sound. For comparison purposes, 30 human participants completed an identical task with the same sounds, objects, and training procedure. The dolphin's ability to correctly match objects to sounds was robust to changes in amplitude with only a minor decrement in performance for short durations. The dolphin failed to recognize sounds that were frequency transposed by plus or minus ½ octaves. Human participants demonstrated robust recognition with all acoustic transformations. The results indicate that this dolphin's acoustic recognition of whistle-like sounds was constrained by absolute pitch. Unlike human speech, which varies considerably in average frequency, signature whistles are relatively stable in frequency, which may have selected for a whistle recognition system invariant to frequency transposition.

Hearing Mechanisms and Noise Metrics Related to Auditory Masking in Bottlenose Dolphins (Tursiops truncatus).

Odontocete cetaceans are acoustic specialists that depend on sound to hunt, forage, navigate, detect predators, and communicate. Auditory masking from natural and anthropogenic sound sources may adversely affect these fitness-related capabilities. The ability to detect a tone in a broad range of natural, anthropogenic, and synthesized noise was tested with bottlenose dolphins using a psychophysical, band-widening procedure. Diverging masking patterns were found for noise bandwidths greater than the width of an auditory filter. Despite different noise types having equal-pressure spectral-density levels (95 dB re 1 µPa(2)/Hz), masked detection threshold differences were as large as 22 dB. Consecutive experiments indicated that noise types with increased levels of amplitude modulation resulted in comodulation masking release due to within-channel and across-channel auditory mechanisms. The degree to which noise types were comodulated (comodulation index) was assessed by calculating the magnitude-squared coherence between the temporal envelope from an auditory filter centered on the signal and temporal envelopes from flanking filters. Statistical models indicate that masked thresholds in a variety of noise types, at a variety of levels, can be explained with metrics related to the comodulation index in addition to the pressure spectral-density level of noise. This study suggests that predicting auditory masking from ocean noise sources depends on both spectral and temporal properties of the noise.

Auditory Effects of Multiple Impulses from a Seismic Air Gun on Bottlenose Dolphins (Tursiops truncatus).

Auditory thresholds were measured in three bottlenose dolphins before and after exposure to ten impulses from a seismic air gun. Thresholds were measured using behavioral and electrophysiological methods to determine the amount of temporary threshold shift induced. The results suggest that the potential for seismic surveys using air guns to cause auditory effects on dolphins may be lower than previously predicted; however, two of the three dolphins exhibited "anticipatory" behavioral changes at the highest exposure condition that suggested they were attempting to mitigate the effects of the exposures.

Effects of multiple impulses from a seismic air gun on bottlenose dolphin hearing and behavior.

To investigate the auditory effects of multiple underwater impulses, hearing thresholds were measured in three bottlenose dolphins before and after exposure to 10 impulses produced by a seismic air gun. Thresholds were measured at multiple frequencies using both psychophysical and electrophysiological (auditory evoked potential) methods. Exposures began at relatively low levels and gradually increased over a period of several months. The highest exposures featured peak sound pressure levels from 196 to 210 dB re 1 µPa, peak-peak sound pressure levels of 200-212 dB re 1 µPa, and cumulative (unweighted) sound exposure levels from 193 to 195 dB re 1 µPa(2)s. At the cessation of the study, no significant increases were observed in psychophysical thresholds; however, a small (9 dB) shift in mean auditory evoked potential thresholds, accompanied by a suppression of the evoked potential amplitude function, was seen in one subject at 8 kHz. At the highest exposure condition, two of the dolphins also exhibited behavioral reactions indicating that they were capable of anticipating and potentially mitigating the effects of impulsive sounds presented at fixed time intervals.