Brian K. Branstetter

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.

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.

Hearing in cetaceans: from natural history to experimental biology.

Sound is a primary sensory cue for most marine mammals, and this is especially true for cetaceans. To passively and actively acquire information about their environment, cetaceans have some of the most derived ears of all mammals, capable of sophisticated, sensitive hearing and auditory processing. These capabilities have developed for survival in an underwater world where sound travels five times faster than in air, and where light is quickly attenuated and often limited at depth, at night, and in murky waters. Cetacean auditory evolution has capitalized on the ubiquity of sound cues and the efficiency of underwater acoustic communication. The sense of hearing is central to cetacean sensory ecology, enabling vital behaviours such as locating prey, detecting predators, identifying conspecifics, and navigating. Increasing levels of anthropogenic ocean noise appears to influence many of these activities. Here, we describe the historical progress of investigations on cetacean hearing, with a particular focus on odontocetes and recent advancements. While this broad topic has been studied for several centuries, new technologies in the past two decades have been leveraged to improve our understanding of a wide range of taxa, including some of the most elusive species. This chapter addresses topics including how sounds are received, what sounds are detected, hearing mechanisms for complex acoustic scenes, recent anatomical and physiological studies, the potential impacts of noise, and mysticete hearing. We conclude by identifying emerging research topics and areas which require greater focus.

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.

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 animals 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.

Dolphin biosonar signals measured at extreme off-axis angles: Insights to sound propagation in the head

Biosonar signals radiated along the beam axis of an Atlantic bottlenose dolphin resemble short transient oscillations. As the azimuth of the measuring hydrophones in the horizontal plane progressively increases with respect to the beam axis the signals become progressively distorted. At approximately ±45°, the signals begin to divide into two components with the time difference between the components increasing with increasing angles. At ±90° or normal to the longitudinal axis of the animal, the time difference between the two pulses measured by the hydrophone on the right side of the dolphins head is, on average, ∼11.9 μs larger than the time differences observed by the hydrophone on the left side of the dolphins head. The center frequency of the first pulse is generally lower, by 33-47 kHz, than the center frequency of the second pulse. When considering the relative locations of the two phonic lips, the data suggest that the signals are being produced by one of the phonic lips and the second pulse resulting from a reflection within the head of the animal. The generation of biosonar signals is a complex process and the propagation pathways through the dolphins head are not well understood.

Auditory masking patterns in bottlenose dolphins (Tursiops truncatus) with natural, anthropogenic, and synthesized noise.

Auditory masking occurs when one sound (usually called noise) interferes with the detection, discrimination, or recognition of another sound (usually called the signal). This interference can lead to detriments in a listeners ability to communicate, forage, and navigate. Most studies of auditory masking in marine mammals have been limited to detection thresholds of pure tones in Gaussian noise. Environmental noise marine mammals encounter is often more complex. In the current study, detection thresholds were estimated for bottlenose dolphins with a 10 kHz signal masked by natural, anthropogenic, and synthesized noise. Using a band-widening paradigm, detection thresholds exhibited a pattern where signal thresholds increased proportionally to bandwidth for narrow band noise. However, when noise bandwidth was greater than a critical band, masking patterns diverged. Subsequent experiments demonstrated that the auditory mechanisms responsible for the divergent masking patterns were related to across-channel comparison and within-valley listening.

Dolphins Can Maintain Vigilant Behavior through Echolocation for 15 Days without Interruption or Cognitive Impairment

In dolphins, natural selection has developed unihemispheric sleep where alternating hemispheres of their brain stay awake. This allows dolphins to maintain consciousness in response to respiratory demands of the ocean. Unihemispheric sleep may also allow dolphins to maintain vigilant states over long periods of time. Because of the relatively poor visibility in the ocean, dolphins use echolocation to interrogate their environment. During echolocation, dolphin produce clicks and listen to returning echoes to determine the location and identity of objects. The extent to which individual dolphins are able to maintain continuous vigilance through this active sense is unknown. Here we show that dolphins may continuously echolocate and accurately report the presence of targets for at least 15 days without interruption. During a total of three sessions, each lasting five days, two dolphins maintained echolocation behaviors while successfully detecting and reporting targets. Overall performance was between 75 to 86% correct for one dolphin and 97 to 99% correct for a second dolphin. Both animals demonstrated diel patterns in echolocation behavior. A 15-day testing session with one dolphin resulted in near perfect performance with no significant decrement over time. Our results demonstrate that dolphins can continuously monitor their environment and maintain long-term vigilant behavior through echolocation.

High-resolution measurement of a bottlenose dolphin's (Tursiops truncatus) biosonar transmission beam pattern in the horizontal plane

Previous measurements of toothed whale echolocation transmission beam patterns have utilized few hydrophones and have therefore been limited to fine angular resolution only near the principal axis or poor resolution over larger azimuthal ranges. In this study, a circular, horizontal planar array of 35 hydrophones was used to measure a dolphins transmission beam pattern with 5° to 10° resolution at azimuths from -150° to +150°. Beam patterns and directivity indices were calculated from both the peak-peak sound pressure and the energy flux density. The emitted pulse became smaller in amplitude and progressively distorted as it was recorded farther off the principal axis. Beyond ±30° to 40°, the off-axis signal consisted of two distinct pulses whose difference in time of arrival increased with the absolute value of the azimuthal angle. A simple model suggests that the second pulse is best explained as a reflection from internal structures in the dolphins head, and does not implicate the use of a second sound source. Click energy was also more directional at the higher source levels utilized at longer ranges, where the center frequency was elevated compared to that of the lower amplitude clicks used at shorter range.

Representing multiple discrimination cues in a computational model of the bottlenose dolphin auditory system

A computational model of the dolphin auditory system was developed to describe how multiple discrimination cues may be represented and employed during echolocation discrimination tasks. The model consisted of a bank of gammatone filters followed by half-wave rectification and low pass filtering. The output of the model resembles a spectrogram; however, the model reflects temporal and spectral resolving properties of the dolphin auditory system. Model outputs were organized to represent discrimination cues related to spectral, temporal and intensity information. Two empirical experiments, a phase discrimination experiment [Johnson et al., Animal Sonar Processes and Performance (Plenum, New York, 1988)] and a cylinder wall thickness discrimination tasks [Au and Pawolski, J. Comp. Physiol. A 170, 41-47 (1992)] were then simulated. Model performance was compared to dolphin performance. Although multiple discrimination cues were potentially available to the dolphin, simulation results suggest temporal information was used in the former experiment and spectral information in the latter. This models representation of sound provides a more accurate approximation to what the dolphin may be hearing compared to conventional spectrograms, time-amplitude, or spectral representations.