Marlee Breese

Behavioral and auditory evoked potential audiograms of a false killer whale (Pseudorca crassidens)

Behavioral and auditory evoked potential (AEP) audiograms of a false killer whale were measured using the same subject and experimental conditions. The objective was to compare and assess the correspondence of auditory thresholds collected by behavioral and electrophysiological techniques. Behavioral audiograms used 3-s pure-tone stimuli from 4 to 45 kHz, and were conducted with a go/no-go modified staircase procedure. AEP audiograms used 20-ms sinusoidally amplitude-modulated tone bursts from 4 to 45 kHz, and the electrophysiological responses were received through gold disc electrodes in rubber suction cups. The behavioral data were reliable and repeatable, with the region of best sensitivity between 16 and 24 kHz and peak sensitivity at 20 kHz. The AEP audiograms produced thresholds that were also consistent over time, with range of best sensitivity from 16 to 22.5 kHz and peak sensitivity at 22.5 kHz. Behavioral thresholds were always lower than AEP thresholds. However, AEP audiograms were completed in a shorter amount of time with minimum participation from the animal. These data indicated that behavioral and AEP techniques can be used successfully and interchangeably to measure cetacean hearing sensitivity.

Predicting temporary threshold shifts in a bottlenose dolphin (Tursiops truncatus): The effects of noise level and duration

Noise levels in the ocean are increasing and are expected to affect marine mammals. To examine the auditory effects of noise on odontocetes, a bottlenose dolphin (Tursiops truncatus) was exposed to octave-band noise (4-8 kHz) of varying durations (<2-30 min) and sound pressures (130-178 dB re 1 microPa). Temporary threshold shift (TTS) occurrence was quantified in an effort to (i) determine the sound exposure levels (SELs) (dB re 1 microPa(2) s) that induce TTS and (ii) develop a model to predict TTS onset. Hearing thresholds were measured using auditory evoked potentials. If SEL was kept constant, significant shifts were induced by longer duration exposures but not for shorter exposures. Higher SELs were required to induce shifts in shorter duration exposures. The results did not support an equal-energy model to predict TTS onset. Rather, a logarithmic algorithm, which increased in sound energy as exposure duration decreased, was a better predictor of TTS. Recovery to baseline hearing thresholds was also logarithmic (approximately -1.8 dB/doubling of time) but indicated variability including faster recovery rates after greater shifts and longer recoveries necessary after longer duration exposures. The data reflected the complexity of TTS in mammals that should be taken into account when predicting odontocete TTS.

The interaction of outgoing echolocation pulses and echoes in the false killer whale’s auditory system: Evoked-potential study

Brain auditory evoked potentials (AEP) associated with echolocation were recorded in a false killer whale Pseudorca crassidens trained to accept suction-cup EEG electrodes and to detect targets by echolocation. AEP collection was triggered by echolocation pulses transmitted by the animal. The target was a hollow aluminum cylinder of strength of -22 dB at a distance from 1 to 8 m. Each AEP record was obtained by averaging more than 1000 individual records. All the records contained two AEP sets: the first one of a constant latency and a second one with a delay proportional to the distance. The timing of these two AEP sets was interpreted as responses to the transmitted echolocation pulse and echo, respectively. The echo-related AEP, although slightly smaller, was comparable to the outgoing click-related AEP in amplitude, even though at a target distance as far as 8 m the echo intensity was as low as -64 dB relative to the transmitted pulse in front of the head. The amplitude of the echo-related AEP was almost independent of distance, even though variation of target distance from 1 to 8 m influenced the echo intensity by as much as 36 dB.

Invariance of evoked-potential echo-responses to target strength and distance in an echolocating false killer whale

Brain auditory evoked potentials (AEPs) were recorded in a false killer whale Pseudorca crassidens trained to accept suction-cup EEG electrodes and to detect targets by echolocation. AEP collection was triggered by echolocation pulses transmitted by the animal. The target strength varied from -22 to -40 dB; the distance varied from 1.5 to 6 m. All the records contained two AEP sets: the first one of a constant latency (transmission-related AEP) and a second one with a delay proportional to the distance (echo-related AEP). The amplitude of echo-related AEPs was almost independent of both target strength and distance, though combined variation of these two parameters resulted in echo intensity variation within a range of 42 dB. The amplitude of transmission-related AEPs was independent of distance but dependent on target strength: the less the target strength, the higher the amplitude. Recording of transmitted pulses has not shown their intensity dependence on target strength. It is supposed that the constancy of echo-related AEP results from variation of hearing sensitivity depending on the target strength and release of echo-related responses from masking by transmitted pulses depending on the distance.

Hearing sensitivity during target presence and absence while a whale echolocates

Hearing sensitivity was measured in a false killer whale during echolocation. Sensitivity was measured using probe stimuli as sinusoidally amplitude modulated signals with a 22.5-kHz carrier frequency and recording auditory evoked potentials as envelope-following responses. The probes were presented and responses were recorded during short 2-s periods when the animal echolocated to detect the presence or absence of a target in a go/no-go paradigm. In the target-absent trials, a hearing threshold of 90.4 dB re 1 muPa was found; in the target-present trials, the threshold was 109.8 dB. Thus, a 19.4-dB difference was found between thresholds in the target-present and target-absent trials. To check the possibility that this difference was the result of different masking degree of the probe by the emitted sonar clicks, click statistics were investigated in similar trials. No indication was found that the energy of the emitted clicks was higher in the target-present than in target-absent trials; on the contrary, mean click level, mean number of clicks per train, and overall train energy was slightly higher in the target-absent trials. Thus the data indicate that the hearing sensitivity of the whale varied depending on target presence or absence.

Active echolocation beam focusing in the false killer whale, Pseudorca crassidens

SUMMARY The odontocete sound production system is highly complex and produces intense, directional signals that are thought to be focused by the melon and the air sacs. Because odontocete echolocation signals are variable and the emitted click frequency greatly affects the echolocation beam shape, investigations of beam focusing must account for frequency-related beam changes. In this study we tested whether the echolocation beam of a false killer whale changed depending on target difficulty and distance while also accounting for frequency-related changes in the echolocation beam. The data indicate that the false killer whale changes its beam size according to target distance and difficulty, which may be a strategy of maximizing the energy of the target echo. We propose that the animal is using a strategy of changing the focal region according to target distance and that this strategy is under active control.

Dolphin hearing during echolocation: evoked potential responses in an Atlantic bottlenose dolphin (Tursiops truncatus)

SUMMARY Auditory evoked potential (AEP) responses were recorded during echolocation in an Atlantic bottlenose dolphin (Tursiops truncatus) trained to accept suction-cup EEG electrodes and detect targets by echolocation. AEP recording was triggered by the echolocation clicks of the animal. Three targets with target strengths of –34, –28 and –22 dB were used at a target distance of 2 to 6.5 m for each target. The results demonstrated that the AEP appeared to both outgoing echolocation clicks and echoes during echolocation, with AEP complexes consisting of alternative positive and negative waves. The echo-related AEP amplitudes were obviously lower than the outgoing click-related AEP amplitudes for all the targets at the investigated target distances. However, for targets with target strengths of –22 and –28 dB, the peak-to-peak amplitudes of the echo-related AEPs were dependent on the target distances. The echo-related AEP response amplitudes increased at further target distances, demonstrating an overcompensation of echo attenuation with target distance in the echo-perception system of the dolphin biosonar. Measurement and analysis of outgoing click intensities showed that the click levels increased with target distance (R) by a factor of approximately 10 to 17.5 logR depending on target strength. The results demonstrated that a dual-component biosonar control system formed by intensity compensation behavior in both the transmission and receiving phases of a biosonar cycle exists synchronously in the dolphin biosonar system.

Evoked-potential recovery during double click stimulation in a whale: A possibility of biosonar automatic gain control

False killer whale Pseudorca crassidens auditory brainstem responses (ABR) were recorded using a double-click stimulation paradigm specifically measuring the recovery of the second response (to the test click) as a function of the inter-click interval (ICI) at various levels of the conditioning and test click. At all click intensities, the slopes of recovery functions were almost constant: 0.6-0.8 microV per ICI decade. Therefore, even when the conditioning-to-test-click level ratio was kept constant, the duration of recovery was intensity-dependent: The higher intensity the longer the recovery. The conditioning-to-test-click level ratio strongly influenced the recovery time: The higher the ratio, the longer the recovery. The dependence was almost linear using a logarithmic ICI scale with a rate of 25-30 dB per ICI decade. These data were used for modeling the interaction between the emitted click and the echo during echolocation, assuming that the two clicks simulated the transmitted and echo clicks. This simulation showed that partial masking of the echo by the preceding emitted click may explain the independence of echo-response amplitude of target distance. However, the distance range where this mechanism is effective depends on the emitted click level: The higher the level, the greater the range. @ 2007 Acoustical Society of America.

Source-to-sensation level ratio of transmitted biosonar pulses in an echolocating false killer whale.

Transmitted biosonar pulses, and the brain auditory evoked potentials (AEPs) associated with those pulses, were synchronously recorded in a false killer whale Pseudorca crassidens trained to accept suction-cup EEG electrodes and to detect targets by echolocation. AEP amplitude was investigated as a function of the transmitted biosonar pulse source level. For that, a few thousand of the individual AEP records were sorted according to the spontaneously varied amplitude of synchronously recorded biosonar pulses. In each of the sorting bins (in 5-dB steps) AEP records were averaged to extract AEP from noise; AEP amplitude was plotted as a function of the biosonar pulse source level. For comparison, AEPs were recorded to external (in free field) sound pulses of a waveform and spectrum similar to those of the biosonar pulses; amplitude of these AEPs was plotted as a function of sound pressure level. A comparison of these two functions has shown that, depending on the presence or absence of a target, the sensitivity of the whales hearing to its own transmitted biosonar pulses was 30 to 45 dB lower than might be expected in a free acoustic field.

Forward masking as a mechanism of automatic gain control in odontocete biosonar: A psychophysical study

In a false killer whale Pseudorca crassidens, echo perception thresholds were measured using a go/no-go psychophysical paradigm and one-up-one-down staircase procedure. Computer controlled echoes were electronically synthesized pulses that were played back through a transducer and triggered by whale emitted biosonar pulses. The echo amplitudes were proportional to biosonar pulse amplitudes; echo levels were specified in terms of the attenuation of the echo sound pressure level near the animals head relative to the source level of the biosonar pulses. With increasing echo delay, the thresholds (echo attenuation factor) decreased from -49.3 dB at 2 ms to -79.5 dB at 16 ms, with a regression slope of -9.5 dB per delay doubling (-31.5 dB per delay decade). At the longer delays, the threshold remained nearly constant around -80.4 dB. Levels of emitted pulses slightly increased with delay prolongation (threshold decrease), with a regression slope of 3.2 dB per delay doubling (10.7 dB per delay decade). The echo threshold dependence on delay is interpreted as a release from forward masking by the preceding emitted pulse. This release may compensate for the echo level decrease with distance, thus keeping the echo sensation level for the animal near constant within a certain distance range.