Killer whales (Orcinus orca) produce ultrasonic whistles

This study reports that killer whales, the largest dolphin, produce whistles with the highest fundamental frequencies ever reported in a delphinid. Using wide-band acoustic sampling from both animal-attached (Dtag) and remotely deployed hydrophone arrays, ultrasonic whistles were detected in three Northeast Atlantic populations but not in two Northeast Pacific populations. These results are inconsistent with analyses suggesting a correlation of maximum frequency of whistles with body size in delphinids, indicate substantial intraspecific variation in whistle production in killer whales, and highlight the importance of appropriate acoustic sampling techniques when conducting comparative analyses of sound repertoires.     

Measuring hearing in the harbor seal (Phoca vitulina): comparison of behavioral and auditory brainstem response techniques

Auditory brainstem response (ABR) and standard behavioral methods were compared by measuring in-air audiograms for an adult female harbor seal (Phoca vitulina). Behavioral audiograms were obtained using two techniques: the method of constant stimuli and the staircase method. Sensitivity was tested from 0.250 to 30 kHz. The seal showed good sensitivity from 6 to 12 kHz [best sensitivity 8.1 dB (re 20 microPa2 x s) RMS at 8 kHz]. The staircase method yielded thresholds that were lower by 10 dB on average than the method of constant stimuli. ABRs were recorded at 2, 4, 8, 16, and 22 kHz and showed a similar best range (8-16 kHz). ABR thresholds averaged 5.7 dB higher than behavioral thresholds at 2, 4, and 8 kHz. ABRs were at least 7 dB lower at 16 kHz, and approximately 3 dB higher at 22 kHz. The better sensitivity of ABRs at higher frequencies could have reflected differences in the seal's behavior during ABR testing and/or bandwidth characteristics of test stimuli. These results agree with comparisons of ABR and behavioral methods performed in other recent studies and indicate that ABR methods represent a good alternative for estimating hearing range and sensitivity in pinnipeds, particularly when time is a critical factor and animals are untrained.     

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.     

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.     

Echolocation clicks from killer whales (Orcinus orca) feeding on herring (Clupea harengus)

Echolocation clicks from Norwegian killer whales feeding on herring schools were recorded using a four-hydrophone array. The clicks had broadband bimodal frequency spectra with low and high frequency peaks at 24 and 108 kHz, respectively. The -10 dB bandwidth was 35 kHz. The average source level varied from 173 to 202 dB re 1 microPa (peak-to-peak) at 1 m. This is considerably lower than source levels described for Canadian killer whales foraging on salmon. It is suggested that biosonar clicks of Norwegian killer whales are adapted for localization of prey with high target strength and acute hearing abilities.     

High-frequency modulated signals of killer whales (Orcinus orca) in the North Pacific

Killer whales in the North Pacific, similar to Atlantic populations, produce high-frequency modulated signals, based on acoustic recordings from ship-based hydrophone arrays and autonomous recorders at multiple locations. The median peak frequency of these signals ranged from 19.6-36.1 kHz and median duration ranged from 50-163 ms. Source levels were 185-193 dB peak-to-peak re: 1 μPa at 1 m. These uniform, repetitive, down-swept signals are similar to bat echolocation signals and possibly could have echolocation functionality. A large geographic range of occurrence suggests that different killer whale ecotypes may utilize these signals.     

The effects of sensory hearing loss on cochlear filter times estimated from auditory brainstem response latencies.

Derived-band auditory brainstem responses (ABRs) were obtained in 43 normal-hearing and 80 cochlear hearing-impaired individuals using clicks and high-pass noise masking. The response times across the cochlea [the latency difference between wave V's of the 5.7- and 1.4-kHz center frequency (CF) derived bands] were calculated for five levels of click stimulation ranging from 53 to 93 dB p.-p.e. SPL (23 to 63 dB nHL) in 10-dB steps. Cochlear response times appeared to shorten significantly with hearing loss, especially when the average pure tone (1 to 8 kHz) hearing loss exceeded 30 dB. Examination of derived-band latencies indicates that this shortening is due to a dramatic decrease of wave V latency in the lower CF derived band. Estimates of cochlear filter times in terms of the number of periods to maximum response (Nmax) were calculated from derived-band latencies corrected for gender-dependent cochlear transport and neural conduction times. Nmax decreased as a function of hearing loss, especially for the low CF derived bands. The functions were similar for both males and females. These results are consistent with broader cochlear tuning due to peripheral hearing loss. Estimating filter response times from ABR latencies enhances objective noninvasive diagnosis and allows delineation of the differential effects of pathology on the underlying cochlear mechanisms involved in cochlear transport and filter build-up times.     

Echolocation signals of free-ranging killer whales (Orcinus orca) and modeling of foraging for chinook salmon (Oncorhynchus tshawytscha)

Fish-eating "resident"-type killer whales (Orcinus orca) that frequent the coastal waters off northeastern Vancouver Island, Canada have a strong preference for chinook salmon (Oncorhynchus tshawytscha). The whales in this region often forage along steep cliffs that extend into the water, echolocating their prey. Echolocation signals of resident killer whales were measured with a four-hydrophone symmetrical star array and the signals were simultaneously digitized at a sample rate of 500 kHz using a lunch-box PC. A portable VCR recorded the images from an underwater camera located adjacent to the array center. Only signals emanating from close to the beam axis (1185 total) were chosen for a detailed analysis. Killer whales project very broadband echolocation signals (Q equal 0.9 to 1.4) that tend to have bimodal frequency structure. Ninety-seven percent of the signals had center frequencies between 45 and 80 kHz with bandwidths between 35 and 50 kHz. The peak-to-peak source level of the echolocation signals decreased as a function of the one-way transmission loss to the array. Source levels varied between 195 and 224 dB re: 1 microPa. Using a model of target strength for chinook salmon, the echo levels from the echolocation signals are estimated for different horizontal ranges between a whale and a salmon. At a horizontal range of 100 m, the echo level should exceed an Orcinus hearing threshold at 50 kHz by over 29 dB and should be greater than sea state 4 noise by at least 9 dB. In moderately heavy rain conditions, the detection range will be reduced substantially and the echo level at a horizontal range of 40 m would be close to the level of the rain noise.     

Pure tone audiograms and possible aminoglycoside-induced hearing loss in belugas (Delphinapterus leucas)

A behavioral response paradigm was used to measure pure-tone hearing sensitivities in two belugas (Delphinapterus leucas). Tests were conducted over a 20-month period at the Point Defiance Zoo and Aquarium, in Tacoma, WA. Subjects were two males, aged 8-10 and 9-11 during the course of the study. Subjects were born in an oceanarium and had been housed together for all of their lives. Hearing thresholds were measured using a modified up/down staircase procedure and acoustic response paradigm where subjects were trained to produce audible responses to test tones and to remain quiet otherwise. Test frequencies ranged from approximately 2 to 130 kHz. Best sensitivities ranged from approximately 40 to 50 dB re 1 microPa at 50-80 kHz and 30-35 kHz for the two subjects. Although both subjects possessed traditional "U-shaped" mammalian audiograms, one subject exhibited significant high-frequency hearing loss above 37 kHz compared to previously published data for belugas. Hearing loss in this subject was estimated to approach 90 dB for frequencies above 50 kHz. Similar ages, ancestry, and environmental conditions between subjects, but a history of ototoxic drug administration in only one subject, suggest that the observed hearing loss was a result of the aminoglycoside antibiotic amikacin.     

Comparison of behavioral and auditory brainstem response measures of threshold shift in rats exposed to loud sound

The purpose of this study was to determine how closely the auditory brainstem response (ABR) can estimate sensorineural threshold shifts in rats exposed to loud sound. Behavioral and ABR thresholds were obtained for tones or noise before and after exposure to loud sound. The results showed that the ABR threshold shift obtained with tone pips estimated the initial pure-tone threshold shifts to within +/-5 dB 11% of the time and the permanent pure-tone threshold shifts 55% of the time, both with large errors. Determining behavioral thresholds for the same tone pips used for the ABR did not improve the agreement between the measures. In contrast, the ABR obtained with octave noise estimated the initial threshold shifts for that noise to within +/-5 dB 25% of the time and the permanent threshold shifts 89% of the time, with much smaller errors. Thus, it appears that the noise-evoked ABR is more accurate in estimating threshold shift than the tone-evoked ABR.     

Masked auditory thresholds in three species of birds, as measured by the auditory brainstem response (L)

Auditory brainstem responses (ABRs) were recorded in adult budgerigars, canaries, and zebra finches in quiet and in three levels of white noise for tone stimuli between 1 and 4 kHz. Similar to behavioral results, masked ABR thresholds increased linearly with increasing noise levels. When the three species are considered together, ABR-derived CRs were higher than behavioral CRs by 18-23 dB between 2 and 4 kHz and by about 30 dB at 1 kHz. This study clarifies the utility of using ABRs for estimating masked auditory thresholds in natural environmental noises in species that cannot be tested behaviorally.     

Rise-fall time effects on the brainstem auditory evoked response: mechanisms

Experiments were conducted to assess the contribution of place mechanisms to the effect of rise--fall time on wave V of the human brainstem auditory evoked response (BAER). Noise bursts of 4- and 10-ms duration were presented at various rise-fall times (0, 1, 2, and 5 ms). Subtractive high-pass masking techniques were used to determine the effect of rise time as a function of derived-band frequency. In general, increasing rise time prolonged wave V latency but did not affect amplitude. Rise-time effects did not depend on derived-band frequency and similar effects were seen in the unmasked conditions. In addition, narrowing the derived band did not alter the observed effects on latency and amplitude. Signal envelope showed no effects on traveling wave velocity. These results suggest that place mechanisms contribute little to changes in the BAER associated with rise--fall time.     

Characteristics of whistles from the acoustic repertoire of resident killer whales (Orcinus orca) off Vancouver Island, British Columbia

The acoustic repertoire of killer whales (Orcinus orca) consists of pulsed calls and tonal sounds, called whistles. Although previous studies gave information on whistle parameters, no study has presented a detailed quantitative characterization of whistles from wild killer whales. Thus an interpretation of possible functions of whistles in killer whale underwater communication has been impossible so far. In this study acoustic parameters of whistles from groups of individually known killer whales were measured. Observations in the field indicate that whistles are close-range signals. The majority of whistles (90%) were tones with several harmonics with the main energy concentrated in the fundamental. The remainder were tones with enhanced second or higher harmonics and tones without harmonics. Whistles had an average bandwidth of 4.5 kHz, an average dominant frequency of 8.3 kHz, and an average duration of 1.8 s. The number of frequency modulations per whistle ranged between 0 and 71. The study indicates that whistles in wild killer whales serve a different function than whistles of other delphinids. Their structure makes whistles of killer whales suitable to function as close-range motivational sounds.     

Correlation among thresholds of evoked otoacoustic emissions,behavioral responses and auditory brainstem responses

In 35 adult human subjects(58 ears)thresholds of rarefactlon clickevoked otoacoustic emission(EOAE),behavioral response(BR)and auditorybrainstem response(ABR)were measured and compared,and correlationcoefficients(r)among them calculated.The results revealed that 86％ of themeasured thresholds were in the range from 10 to 45 dB(nHL)for EOAE,88％ from 10 to 25 dB for BR and 88％ from 10 to 35 for ABR.The correlationcoefficients for the whole threshold sample were 0.415(p<0.002)for EOAE vs.BR,0.501(p<0.001)for EOAE vs.ABR and 0.702(p<0.001)for ABR vs.BR,all indicating highly significant correlation.However,for those ears whoseBR,ABR or EOAE thresholds were elevated,equaling to or exceeding 25 dB,there is no significant correlation between thresholds of EOAE and BR and ofEOAE and ABR(r range:0.176-0.310,p>0.05).In contrast,significantcorrelation between BR and ABR thresholds in the same conditions still re-mained(r:0.533-0.720,p<0.05).The experimental results indicate that thecorrelation between EOAE and hea     

California sea lion (Zalophus californianus) aerial hearing sensitivity measured using auditory steady-state response and psychophysical methods

Although electrophysiological methods of measuring the hearing sensitivity of pinnipeds are not yet as refined as those for dolphins and porpoises, they appear to be a promising supplement to traditional psychophysical procedures. In order to further standardize electrophysiological methods with pinnipeds, a within-subject comparison of psychophysical and auditory steady-state response (ASSR) measures of aerial hearing sensitivity was conducted with a 1.5-yr-old California sea lion. The psychophysical audiogram was similar to those previously reported for otariids, with a U-shape, and thresholds near 10 dB re 20 μPa at 8 and 16 kHz. ASSR thresholds measured using both single and multiple simultaneous amplitude-modulated tones closely reproduced the psychophysical audiogram, although the mean ASSR thresholds were elevated relative to psychophysical thresholds. Differences between psychophysical and ASSR thresholds were greatest at the low- and high-frequency ends of the audiogram. Thresholds measured using the multiple ASSR method were not different from those measured using the single ASSR method. The multiple ASSR method was more rapid than the single ASSR method, and allowed for threshold measurements at seven frequencies in less than 20 min. The multiple ASSR method may be especially advantageous for hearing sensitivity measurements with otariid subjects that are untrained for psychophysical procedures.     

Frequency- and level-dependent changes in auditory brainstem responses (ABRS) in developing mice

The development of the auditory brainstem response was studied to quantitatively assess its dependence on stimulus frequency and level. Responses were not observed to stimuli > or =16 kHz on P12, however, the full range of responsive frequencies included in the study was observed by P14. Response thresholds were high on P12, exceeding 100 dB SPL for all stimuli tested. The rate of threshold development increased progressively for stimulus frequencies between -2 and 10 kHz, with the most rapid changes occurring at frequencies >10 kHz. Adultlike thresholds were observed by P18. Response latencies and interpeak intervals matured rapidly over the course of the second and third postnatal weeks and did not achieve adultlike characteristics until after P18. Latencies of higher-order peaks were progressively and sequentially delayed relative to wave I. Wave I amplitudes developed nonmonotonically, growing during the first 24 days and stabilizing at adult values by approximately P36. Slopes of wave I amplitude-and latency-level curves were significantly steeper than those of adults during the neonatal period and the outcome of input-output analyses, as well as frequency-specific maturational profiles, support developmental models in which function initially matures in the mid-frequency range and proceeds, simultaneously, in both apical and basal directions.     

The effect of broadband noise on the human brainstem auditory evoked response. II. Frequency specificity

A series of experiments evaluated the effects of broadband noise (ipsilateral) on wave V of the brainstem auditory evoked response (BAER) elicited by tone bursts or clicks in the presence of high-pass masking noise. Experiment 1 used 1000- and 4000-Hz, 60-dB nHL tone bursts in the presence of broadband noise. With increasing noise level, wave V latency shift was greater for the 1000-Hz tone bursts, while amplitude decrements were similar for both tone-burst frequencies. Experiment 2 varied high-pass masker cutoff frequency and the level of subtotal masking in the presence of 50-dB nHL clicks. The effects of subtotal masking on wave V (increase in latency and decrease in amplitude) increased with increasing derived-band frequency. Experiment 3 covaried high-pass masker cutoff frequency and subtotal masking level for 1000- and 4000-Hz tone-burst stimuli. The effect of subtotal masking on wave V latency was reduced for both tone-burst frequencies when the response-generating region of the cochlear partition was limited by high-pass maskers. The results of these three experiments suggest that most of the wave V latency shift associated with increasing levels of broadband noise is mediated by a place mechanism when the stimulus is a moderate intensity (60 dB nHL), low-frequency (1000 Hz) tone burst. However, the interpretation of the latency shifts produced by broadband noise for 4000-Hz tone-burst stimuli is made more complex by multiple technical factors discussed herein.     

An exact method of regularity analysis for auditory brainstem neurons (L)

The standard regularity analysis for spike trains in cochlear nucleus neurons evoked by tonebursts first proposed by Bourk is widely used, primarily as one of the criteria for classification of such neurons. It is shown that this procedure does not estimate quite what it is supposed to, and introduces unnecessary noise to its results due to its use of bins. Instead the desired quantities (mean and coefficient of variation of the lengths of all inter-spike intervals in progress as a function of time since stimulus onset) can all be exactly calculated directly from the spike train without the need for data binning. The implications for classification and other studies are discussed.     

The effect of broadband noise on the human brainstem auditory evoked response. I. Rate and intensity effects

A series of experiments investigated the effects of continuous broadband noise (ipsilateral) on wave V of the click-evoked brainstem auditory evoked response (BAER). In general, a broadband noise masker increases the latency and decreases the amplitude of wave V. Varying both click and noise intensity, it was found that noise levels above about 40 dB SPL increase the latency and decrease the amplitude of wave V, regardless of click intensity. The effects of noise on wave V amplitude appear constant across click intensity, whereas the effects of a constant noise level on wave V latency decrease at higher click intensities. Both masking and adaptation increase wave V latency, but their combined effects are occlusive: rate-induced wave V latency shift decreases in the presence of continuous broadband noise. The clinical and theoretical implications of these findings are discussed.