The perception of complex tones by a false killer whale (Pseudorca crassidens)

Complex tonal whistles are frequently produced by some odontocete species. However, no experimental evidence exists regarding the detection of complex tones or the discrimination of harmonic frequencies by a marine mammal. The objectives of this investigation were to examine the ability of a false killer whale to discriminate pure tones from complex tones and to determine the minimum intensity level of a harmonic tone required for the whale to make the discrimination. The study was conducted with a go/no-go modified staircase procedure. The different stimuli were complex tones with a fundamental frequency of 5 kHz with one to five harmonic frequencies. The results from this complex tone discrimination task demonstrated: (1) that the false killer whale was able to discriminate a 5 kHz pure tone from a complex tone with up to five harmonics, and (2) that discrimination thresholds or minimum intensity levels exist for each harmonic combination measured. These results indicate that both frequency level and harmonic content may have contributed to the false killer whale's discrimination of complex tones.     

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.     

Single-lobed frequency-dependent beam shape in an echolocating false killer whale (Pseudorca crassidens)

Recent studies indicate some odontocetes may produce echolocation beams with a dual-lobed vertical structure. The shape of the odontocete echolocation beam was further investigated in a false killer whale performing an echolocation discrimination task. Clicks were recorded with an array of 16 hydrophones and frequency-dependent amplitude plots were constructed to assess beam shape. The majority of the echolocation clicks were single-lobed in structure with most energy located between 20 and 80 kHz. These data indicate the false killer whale does not produce a dual-lobed structure, as has been shown in bottlenose dolphins, which may be a function of lowered frequencies in the emitted signal due to hearing loss.     

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.     

Target detection, shape discrimination, and signal characteristics of an echolocating false killer whale (Pseudorca crassidens).

This study demonstrated the ability of a false killer whale (Pseudorca crassidens) to discriminate between two targets and investigated the parameters of the whale's emitted signals for changes related to test conditions. Target detection performance comparable to the bottlenose dolphin's (Tursiops truncatus) has previously been reported for echolocating false killer whales. No other echolocation capabilities have been reported. A false killer whale, naive to conditioned echolocation tasks, was initially trained to detect a cylinder in a "go/no-go" procedure over ranges of 3 to 8 m. The transition from a detection task to a discrimination task was readily achieved by introducing a spherical comparison target. Finally, the cylinder was successfully compared to spheres of two different sizes and target strengths. Multivariate analyses were used to evaluate the parameters of emitted signals. Duncan's multiple range tests showed significant decreases (df = 185, p less than 0.05) in both source level and bandwidth in the transition from detection to discrimination. Analysis of variance revealed a significant decrease in the number of clicks over test conditions [F(5.26) = 5.23, p less than 0.0001]. These data suggest that the whale relied on cues relevant to target shape as well as target strength, that changes in source level and bandwidth were task-related, that the decrease in clicks was associated with learning experience, and that Pseudorca's ability to discriminate shapes using echolocation may be comparable to that of Tursiops truncatus.     

Underwater psychophysical audiogram of a young male California sea lion (Zalophus californianus)

Auditory evoked potential (AEP) data are commonly obtained in air while sea lions are under gas anesthesia; a procedure that precludes the measurement of underwater hearing sensitivity. This is a substantial limitation considering the importance of underwater hearing data in designing criteria aimed at mitigating the effects of anthropogenic noise exposure. To determine if some aspects of underwater hearing sensitivity can be predicted using rapid aerial AEP methods, this study measured underwater psychophysical thresholds for a young male California sea lion (Zalophus californianus) for which previously published aerial AEP thresholds exist. Underwater thresholds were measured in an aboveground pool at frequencies between 1 and 38 kHz. The underwater audiogram was very similar to those previously published for California sea lions, suggesting that the current and previously obtained psychophysical data are representative for this species. The psychophysical and previously measured AEP audiograms were most similar in terms of high-frequency hearing limit (HFHL), although the underwater HFHL was sharper and occurred at a higher frequency. Aerial AEP methods are useful for predicting reductions in the HFHL that are potentially independent of the testing medium, such as those due to age-related sensorineural hearing loss.     

Similarities in echolocation strategy and click characteristics between a Pseudorca crassidens and a Tursiops truncatus

A previous comparative analysis of normalized click amplitude spectra from a Tursiops truncatus has shown that those frequencies with the lowest click-to-click variability in spectral content were the frequencies the animal paid attention to during target discrimination tasks. In that case, the dolphin only paid attention to the frequency range between 29-42 kHz which had a significantly higher degree of consistency in spectral content than frequencies above 42 kHz. Here it is shown that despite their morphological and behavioral differences, this same pattern of consistency was used by a Pseudorca crassidens performing a similar discrimination task. This comparison between species provides a foundation for using spectral level variability to determine the frequencies most important for echolocation in rare species and non-captive animals. Such results provide key information for successful management.     

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.     

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 whale's hearing to its own transmitted biosonar pulses was 30 to 45 dB lower than might be expected in a free acoustic field.     

Interaction of emitted sonar pulses and simulated echoes in a false killer whale: an evoked-potential study

Auditory evoked potentials (AEP) were recorded during echolocation in a false killer whale Pseudorca crassidens. An electronically synthesized and played-back (simulated) echo was triggered by an emitted biosonar pulse, and its intensity was proportional to that of the emitted click. The delay and transfer factor of the echo relative to the emitted click was controlled by the operator. The echo delay varied from 2 to 16 ms (by two-fold steps), and the transfer factor varied within ranges from -45 to -30 dB at the 2-ms delay to -60 to -45 dB at the 16-ms delay. Echo-related AEPs featured amplitude dependence both on echo delay at a constant transfer factor (the longer the delay, the higher amplitude) and on echo transfer factor at a constant delay (the higher transfer factor, the higher amplitude). Conjunctional variation of the echo transfer factor and delay kept the AEP amplitude constant when the delay to transfer factor trade was from -7.1 to -8.4 dB per delay doubling. The results confirm the hypothesis that partial forward masking of the echoes by the preceding emitted sonar pulses serves as a time-varying automatic gain control in the auditory system of echolocating odontocetes.     

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.     

Free-field audiogram of the Japanese macaque (Macaca fuscata)

The audiograms of three Japanese macaques and seven humans were determined in a free-field environment using loudspeakers. The monkeys and humans were tested using tones ranging from 8 Hz to 40 kHz and 4 Hz to 22.4 kHz, respectively. At a level of 60 dB sound pressure level the monkeys were able to hear tones extending from 28 Hz to 37 kHz with their best sensitivity of 1 dB occurring at 4 kHz. The human 60-dB hearing range extended from 31 Hz to 17.6 kHz with a best sensitivity of -10 dB at 2 and 4 kHz. These results indicate that the Japanese macaque has low-frequency hearing equal to that of humans and better than that indicated by previous audiograms obtained using headphones.     

Quantifying complex patterns of bioacoustic variation: use of a neural network to compare killer whale (Orcinus orca) dialects

A quantitative measure of acoustic similarity is crucial to any study comparing vocalizations of different species, social groups, or individuals. The goal of this study was to develop a method of extracting frequency contours from recordings of pulsed vocalizations and to test a nonlinear index of acoustic similarity based on the error of an artificial neural network at classifying them. Since the performance of neural networks depends on the amount of consistent variation in the training data, this technique can be used to assess such variation from samples of acoustic signals. The frequency contour extraction and the neural network index were tested on samples of one call type shared by nine social groups of killer whales. For comparison, call similarity was judged by three human subjects in pairwise classification tasks. The results showed a significant correlation between the neural network index and the similarity ratings by the subjects. Both measures of acoustic similarity were significantly correlated with the groups' association patterns, indicating that both methods of quantifying acoustic similarity are biologically meaningful. An index based on neural network analysis therefore represents an objective and repeatable means of measuring acoustic similarity, and allows comparison of results across studies, species and time.     

Automatic classification of killer whale vocalizations using dynamic time warping

A set of killer whale sounds from Marineland were recently classified automatically [Brown et al., J. Acoust. Soc. Am. 119, EL34-EL40 (2006)] into call types using dynamic time warping (DTW), multidimensional scaling, and kmeans clustering to give near-perfect agreement with a perceptual classification. Here the effectiveness of four DTW algorithms on a larger and much more challenging set of calls by Northern Resident whales will be examined, with each call consisting of two independently modulated pitch contours and having considerable overlap in contours for several of the perceptual call types. Classification results are given for each of the four algorithms for the low frequency contour (LFC), the high frequency contour (HFC), their derivatives, and weighted sums of the distances corresponding to LFC with HFC, LFC with its derivative, and HFC with its derivative. The best agreement with the perceptual classification was 90% attained by the Sakoe-Chiba algorithm for the low frequency contours alone.     

Mathematics of pulsed vocalizations with application to killer whale biphonation

Formulas for the spectra of pulsed vocalizations for both the continuous and discrete cases are rigorously derived from basic formulas for Fourier analysis, a topic discussed qualitatively in Watkins' classic paper on "the harmonic interval" ["The harmonic interval: Fact or artifact in spectral analysis of pulse trains," in Marine Bioacoustics 2, edited by W. N. Tavogla (Pergamon, New York, 1967), pp. 15-43]. These formulas are summarized in a table for easy reference, along with most of the corresponding graphs. The case of a "pulse tone" is shown to involve multiplication of two temporal wave forms, corresponding to convolution in the frequency domain. This operation is discussed in detail and shown to be equivalent to a simpler approach using a trigonometric formula giving sum and difference frequencies. The presence of a dc component in the temporal wave form, which implies physically that there is a net positive pressure at the source, is discussed, and examples of the corresponding spectra are calculated and shown graphically. These have application to biphonation (two source signals) observed for some killer whale calls and implications for a source mechanism. A MATLAB program for synthesis of a similar signal is discussed and made available online.     

Click production during breathing in a sperm whale (Physeter macrocephalus)

A sperm whale (Physeter macrocephalus) was observed at the surface with above- and underwater video and synchronized underwater sound recordings. During seven instances the whale ventilated its lungs while clicking. From this observation it is inferred that click production is achieved by pressurizing air in the right nasal passage, pneumatically disconnected from the lungs and the left nasal passage, and that air flows anterior through the phonic lips into the distal air sac. The capability of breathing and clicking at the same time is unique among studied odontocetes and relates to the extreme asymmetry of the sperm whale sound-producing forehead.     

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.     

Effects of noise levels and call types on the source levels of killer whale calls

Accurate parameter estimates relevant to the vocal behavior of marine mammals are needed to assess potential effects of anthropogenic sound exposure including how masking noise reduces the active space of sounds used for communication. Information about how these animals modify their vocal behavior in response to noise exposure is also needed for such assessment. Prior studies have reported variations in the source levels of killer whale sounds, and a more recent study reported that killer whales compensate for vessel masking noise by increasing their call amplitude. The objectives of the current study were to investigate the source levels of a variety of call types in southern resident killer whales while also considering background noise level as a likely factor related to call source level variability. The source levels of 763 discrete calls along with corresponding background noise were measured over three summer field seasons in the waters surrounding the San Juan Islands, WA. Both noise level and call type were significant factors on call source levels (1-40 kHz band, range of 135.0-175.7 dB(rms) re 1 [micro sign]Pa at 1 m). These factors should be considered in models that predict how anthropogenic masking noise reduces vocal communication space in marine mammals.     

Underwater auditory localization by a swimming harbor seal (Phoca vitulina)

The underwater sound localization acuity of a swimming harbor seal (Phoca vitulina) was measured in the horizontal plane at 13 different positions. The stimulus was either a double sound (two 6-kHz pure tones lasting 0.5 s separated by an interval of 0.2 s) or a single continuous sound of 1.2 s. Testing was conducted in a 10-m-diam underwater half circle arena with hidden loudspeakers installed at the exterior perimeter. The animal was trained to swim along the diameter of the half circle and to change its course towards the sound source as soon as the signal was given. The seal indicated the sound source by touching its assumed position at the board of the half circle. The deviation of the seals choice from the actual sound source was measured by means of video analysis. In trials with the double sound the seal localized the sound sources with a mean deviation of 2.8 degrees and in trials with the single sound with a mean deviation of 4.5 degrees. In a second experiment minimum audible angles of the stationary animal were found to be 9.8 degrees in front and 9.7 degrees in the back of the seal's head.