Discrimination of phase altered targets by an echolocating Atlantic bottlenose dolphin.

Sensitivity of echolocating dolphins to phase changes within echoes may be a vital piece of information when constructing echolocation models. Previous experiments have yielded ambiguous results leaving it unclear what cues might have been used by passively listening dolphins to discriminate between different phase altered signals. This study used a phantom echo generator to produce computer controlled echoes. The dolphin interacted with the system in a real echolocation task to discriminate between simulated targets that were unaltered and those that had a 180° phase shift. The frequency amplitude spectral content between the two targets was the same. There were no temporal differences between the two targets. The only cue that the dolphin could use to discriminate between them was the 180° phase shift. The dolphin preformed at a success level of 40% in discriminating the two echoes. This indicates that the 180° phase shift was not perceived.

Spatial orientation of different frequencies within the echolocation beam of a Tursiops truncatus and Pseudorca crassidens.

A two-dimensional array of 16 hydrophones was created to map the spatial distribution of different frequencies within the echolocation beam of a Tursiops truncatus and a Pseudorca crassidens. It was previously shown that both the Tursiops and Pseudorca only paid attention to frequencies between 29 and 42 kHz while echolocating. Both individuals tightly focused the 30 kHz frequency and the spatial location of the focus was consistently pointed toward the target. At 50 kHz the beam was less focused and less precisely pointed at the target. At 100 kHz the focus was often completely lost and was not pointed at the target. This indicates that these individuals actively focused the beam toward the target only in the frequency range they paid attention to. Frequencies outside this range were left unfocused and undirected. This focusing was probably achieved through sensorimotor control of the melon morphology and nasal air sacs. This indicates that both morphologically different species can control the spatial distribution of different frequency ranges within the echolocation beam to create consistent ensonation of desired targets.

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

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.

Change in echolocation signals with hearing loss in a false killer whale (Pseudorca crassidens).

The echolocation signals of a false killer whale (Pseudorca crassidens) were collected during a wall thickness discrimination task and compared to clicks recorded during an identical experiment in 1992. During the sixteen year time period, the subject demonstrated a loss of high frequency hearing of about 70 kHz. Clicks between the two experiments were compared to investigate the effect of hearing loss on echolocation signals. There was a significant reduction in the peak frequency, center frequency and source level of clicks between the two time periods. Additionally, the subject currently produces more signals with low frequency peaks and fewer signals with high frequency peaks than she did in 1992. These results indicate the subject changed its echolocation signals to match its range of best hearing.

Functional bandwidth of an echolocating Atlantic bottlenose dolphin (Tursiops truncatus).

The frequency band that an Atlantic bottlenose dolphin (Tursiops truncatus) used to perform an echolocation target discrimination task was determined using computer simulated phantom targets. The dolphin was trained to discriminate frequency filtered phantom targets from unfiltered ones in a go/no-go paradigm. The dolphin's performance indicated perception of echo alteration only when applied filters interfered with the frequency band between 29 and 42 kHz. The dolphin did not behaviorally convey perception of applied filters that affected frequencies outside this functional bandwidth, such as a low pass 43 kHz or a high pass 28 kHz filter. The upper limit of the functional bandwidth at 42 kHz corresponded with the dolphin's upper hearing limit of 45 kHz, as determined through auditory evoked potential measurements. The lower limit of the functional bandwidth corresponded to a drop in intensity below 30 kHz within the dolphin's echolocation clicks. The randomized presentation of different filters showed that the dolphin paid attention to the entire 29-42 kHz band for each trial, not just subsets. The absence of temporal cues between some of the targets the dolphin could discriminate indicated that in these cases the target discrimination cues were based solely on the frequency content.

Phantom echo highlight amplitude and temporal difference resolutions of an echolocating dolphin, Tursiops truncatus.

A dolphin's ability to discriminate targets may depend greatly on the relative amplitudes and the time separations of echo highlights within the received signal. Previous experiments with dolphins varied the physical parameters of targets, but did not fully investigate how changes in these parameters correspond with the scattered acoustic wave forms and the dolphin's subsequent response. This experiment utilizes a phantom echo system to test a dolphin's detection response to relative amplitude differences of secondary and trailing echo highlights and the time separation differences of all the echo highlights both within and outside the animal's integration window. By electronically manipulating the amplitude and temporal separation of the echo highlights, the underlying acoustic classification cues are more efficiently investigated. The animal successfully discriminated between a standard echo signal and one with the secondary highlight amplitude lowered by 7 dB from the standard. Furthermore, the animal successfully discriminated between a standard echo signal and one with the trailing highlight amplitude lowered by 3 dB from the standard and also a standard echo signal and one with a time separation of 150 mus between the secondary and trailing highlights.

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.

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.

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.