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Biomedical subjects

Whitlow W L Au

Publications and source records attributed to Whitlow W L Au.

At least 19 recordsLinked to original sources

Acoustic features of objects matched by an echolocating bottlenose dolphin.

The focus of this study was to investigate how dolphins use acoustic features in returning echolocation signals to discriminate among objects. An echolocating dolphin performed a match-to-sample task with objects that varied in size, shape, material, and texture. After the task was completed, the features of the object echoes were measured (e.g., target strength, peak frequency). The dolphin's error patterns were examined in conjunction with the between-object variation in acoustic features to identify the acoustic features that the dolphin used to discriminate among the objects. The present study explored two hypotheses regarding the way dolphins use acoustic information in echoes: (1) use of a single feature, or (2) use of a linear combination of multiple features. The results suggested that dolphins do not use a single feature across all object sets or a linear combination of six echo features. Five features appeared to be important to the dolphin on four or more sets: the echo spectrum shape, the pattern of changes in target strength and number of highlights as a function of object orientation, and peak and center frequency. These data suggest that dolphins use multiple features and integrate information across echoes from a range of object orientations.

Acoustics↗

Testing the odontocete acoustic prey debilitation hypothesis: no stunning results.

The hypothesis that sounds produced by odontocetes can debilitate fish was examined. The effects of simulated odontocete pulsed signals on three species of fish commonly preyed on by odontocetes were examined, exposing three individuals of each species as well as groups of four fish to a high-frequency click of a bottlenose dolphin [peak frequency (PF) 120 kHz, 213-dB peak-to-peak exposure level (EL)], a midfrequency click modeled after a killer whale's signal (PF 55 kHz, 208-dB EL), and a low-frequency click (PF 18 kHz, 193-dB EL). Fish were held in a 50-cm diameter net enclosure immediately in front of a transducer where their swimming behavior, orientation, and balance were observed with two video cameras. Clicks were presented at constant rates and in graded sweeps simulating a foraging dolphin's "terminal buzz." No measurable change in behavior was observed in any of the fish for any signal type or pulse modulation rate, despite the fact that clicks were at or near the maximum source levels recorded for odontocetes. Based on the results, the hypothesis that acoustic signals of odontocetes alone can disorient or "stun" prey cannot be supported.

Acoustic Stimulation↗

Acoustic properties of humpback whale songs.

A vertical array of five hydrophones was used to measure the acoustic field in the vertical plane of singing humpback whales. Once a singer was located, two swimmers with snorkel gear were deployed to determine the orientation of the whale and position the boat so that the array could be deployed in front of the whale at a minimum standoff distance of at least 10 m. The spacing of the hydrophones was 7 m with the deepest hydrophone deployed at a depth of 35 m. An eight-channel TASCAM recorder with a bandwidth of 24 kHz was used to record the hydrophone signals. The location (distance and depth) of the singer was determined by computing the time of arrival differences between the hydrophone signals. The maximum source level varied between individual units in a song, with values between 151 and 173 dB re 1 microPa. One of the purposes of this study was to estimate potential sound exposure of nearby conspecifics. The acoustic field determined by considering the relative intensity of higher frequency harmonics in the signals indicated that the sounds are projected in the horizontal direction despite the singer being canted head downward anywhere from about 25 degrees to 90 degrees. High-frequency harmonics extended beyond 24 kHz, suggesting that humpback whales may have an upper frequency limit of hearing as high as 24 kHz.

Acoustics↗

Acoustic radiation from the head of echolocating harbor porpoises (Phocoena phocoena).

An experiment was conducted to investigate the sound pressure patterns on the melon of odontocetes by using four broadband hydrophones embedded in suction cups to measure echolocation signals on the surface of the forehead of two harbor porpoises (Phocoena phocoena). It has long been hypothesized that the special lipids found in the melon of odontocetes, and not in any other mammals, focus sounds produced in the nasal region that then propagate through the melon, producing a beam that is directional in both the horizontal and vertical planes. The results of our measurements supported the melon-focusing hypothesis, with the maximum click amplitude, representing the axis of the echolocation beam, located approximately 5.6-6.1 cm from the edge of the animal's upper lip along the midline of the melon. The focusing is not sharp but is sufficient to produce a transmission beam of about 16 degrees. Click amplitude dropped off rapidly at locations away from the location of site of maximum amplitude. Based on comparisons of forehead anatomy from similar sized porpoises, the beam axis coincided with a pathway extending from the phonic lips through the axis of the low-density/low sound velocity lipid core of the melon. The significant interaction between click number and hydrophone position suggests that the echolocation signals can take slightly different pathways through the melon, probably as a result of how the signals are launched by the production mechanism and the position of the acoustically reflective air sacs.

Acoustics↗

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.

Animals↗

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.

Animals↗

Echolocation signals of dusky dolphins (Lagenorhynchus obscurus) in Kaikoura, New Zealand.

An array of four hydrophones arranged in a symmetrical star configuration was used to measure the echolocation signals of the dusky dolphin (Lagenorhynchus obscurus) near the Kaikoura Peninsula, New Zealand. Most of the echolocation signals had bi-modal frequency spectra with a low-frequency peak between 40 and 50 kHz and a high-frequency peak between 80 and 110 kHz. The low-frequency peak was dominant when the source level was low and the high frequency peak dominated when the source level was high. The center frequencies in the dusky broadband echolocation signals are among the highest of dolphins measured in the field. Peak-to-peak source levels as high as 210 dB re 1 microPa were measured, although the average was much lower in value. The levels of the echolocation signals are about 9-12 dB lower than for the larger white-beaked dolphin (Lagenorhynchus albirostris) which belongs to the same genus but is over twice as heavy as the dusky dolphins. The source level varied in amplitude approximately as a function of the one-way transmission loss for signals traveling from the animals to the array. The wave form and spectrum of the echolocation signals were similar to those of other dolphins measured in the field.

Animals↗

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.

Acoustic Stimulation↗

Geographic variations in the whistles of spinner dolphins (Stenella longirostris) of the Main Hawai'ian Islands.

Geographic variations in the whistles of Hawai'ian spinner dolphins are discussed by comparing 27 spinner dolphin pods recorded in waters off the Islands of Kaua'i, O'ahu, Lana'i, and Hawai'i. Three different behavioral states, the number of dolphins observed in each pod, and ten parameters extracted from each whistle contour were considered by using clustering and discriminant function analyses. The results suggest that spinner dolphin pods in the Main Hawai'ian Islands share characteristics in approximately 48% of their whistles. Spinner dolphin pods had similar whistle parameters regardless of the island, location, and date when they were sampled and the dolphins' behavioral state and pod size. The term "whistle-specific subgroup" (WSS) was used to designate whistle groups with similar whistles parameters (which could have been produced in part by the same dolphins). The emission rate of whistles was higher when spinner dolphins were socializing than when they were traveling or resting, suggesting that whistles are mainly used during close-range interactions. Spinner dolphins also seem to vary whistle duration according to their general behavioral state. Whistle duration and the number of turns and steps of a whistle may be more important in delivering information at the individual level than whistle frequency parameters.

Animals↗

Automatic gain control in the echolocation system of dolphins.

In bats and technological sonars, the gain of the receiver is progressively increased with time after the transmission of a signal to compensate for acoustic propagation loss. The current understanding of dolphin echolocation indicates that automatic gain control is not a part of their sonar system. In order to test this understanding, we have performed field measurements of free-ranging echolocating dolphins. Here we show that dolphins do possess an automatic gain control mechanism, but that it is implemented in the transmission phase rather than the receiving phase of a sonar cycle. We find that the amplitude of the dolphins' echolocation signals are highly range dependent; this amplitude increases with increasing target range, R, in a 20 log(R) fashion to compensate for propagation loss. If the echolocation target is a fish school with many sound scatterers, the echoes from the school will remain nearly constant with range as the dolphin closes in on it. This characteristic has the same effect as time-varying gain in bats and technological sonar when considered from a sonar system perspective.

Acoustics↗

Echolocation signals of wild Atlantic spotted dolphin (Stenella frontalis).

An array of four hydrophones arranged in a symmetrical star configuration was used to measure the echolocation signals of the Atlantic spotted dolphin (Stenella frontalis) in the Bahamas. The spacing between the center hydrophone and the other hydrophones was 45.7 cm. A video camera was attached to the array and a video tape recorder was time synchronized with the computer used to digitize the acoustic signals. The echolocation signals had bi-modal frequency spectra with a low-frequency peak between 40 and 50 kHz and a high-frequency peak between 110 and 130 kHz. The low-frequency peak was dominant when the signal the source level was low and the high-frequency peak dominated when the source level was high. Peak-to-peak source levels as high as 210 dB re 1 microPa were measured. The source level varied in amplitude approximately as a function of the one-way transmission loss for signals traveling from the animals to the array. The characteristics of the signals were similar to those of captive Tursiops truncatus, Delphinapterus leucas and Pseudorca crassidens measured in open waters under controlled conditions.

Animals↗

Echolocation in the Risso's dolphin, Grampus griseus.

The Risso's dolphin (Grampus griseus) is an exclusively cephalopod-consuming delphinid with a distinctive vertical indentation along its forehead. To investigate whether or not the species echolocates, a female Risso's dolphin was trained to discriminate an aluminum cylinder from a nylon sphere (experiment 1) or an aluminum sphere (experiment 2) while wearing eyecups and free swimming in an open-water pen in Kaneohe Bay, Hawaii. The dolphin completed the task with little difficulty despite being blindfolded. Clicks emitted by the dolphin were acquired at average amplitudes of 192.6 dB re 1 microPa, with estimated sources levels up to 216 dB re 1 microPa-1 m. Clicks were acquired with peak frequencies as high as 104.7 kHz (Mf(p) = 47.9 kHz), center frequencies as high as 85.7 kHz (Mf(0) = 56.5 kHz), 3-dB bandwidths up to 94.1 kHz (M(BW) = 39.7 kHz), and root-mean-square bandwidths up to 32.8 kHz (M(RMS) = 23.3 kHz). Click durations were between 40 and 70 micros. The data establish that the Risso's dolphin echolocates, and that, aside from slightly lower amplitudes and frequencies, the clicks emitted by the dolphin were similar to those emitted by other echolocating odontocetes. The particular acoustic and behavioral findings in the study are discussed with respect to the possible direction of the sonar transmission beam of the species.

Animals↗

Audiogram of a striped dolphin (Stenella coeruleoalba).

The underwater hearing sensitivity of a striped dolphin was measured in a pool using standard psycho-acoustic techniques. The go/no-go response paradigm and up-down staircase psychometric method were used. Auditory sensitivity was measured by using 12 narrow-band frequency-modulated signals having center frequencies between 0.5 and 160 kHz. The 50% detection threshold was determined for each frequency. The resulting audiogram for this animal was U-shaped, with hearing capabilities from 0.5 to 160 kHz (8 1/3 oct). Maximum sensitivity (42 dB re 1 microPa) occurred at 64 kHz. The range of most sensitive hearing (defined as the frequency range with sensitivities within 10 dB of maximum sensitivity) was from 29 to 123 kHz (approximately 2 oct). The animal's hearing became less sensitive below 32 kHz and above 120 kHz. Sensitivity decreased by about 8 dB per octave below 1 kHz and fell sharply at a rate of about 390 dB per octave above 140 kHz.

Acoustic Stimulation↗

Temporary threshold shifts and recovery following noise exposure in the Atlantic bottlenosed dolphin (Tursiops truncatus).

Behaviorally determined hearing thresholds for a 7.5-kHz tone for an Atlantic bottlenosed dolphin (Tursiops truncatus) were obtained following exposure to fatiguing low-frequency octave band noise. The fatiguing stimulus ranged from 4 to 11 kHz and was gradually increased in intensity to 179 dB re 1 microPa and in duration to 55 min. Exposures occurred no more frequently than once per week. Measured temporary threshold shifts averaged 11 dB. Threshold determination took at least 20 min. Recovery was examined 360, 180, 90, and 45 min following exposure and was essentially complete within 45 min.

Acoustic Stimulation↗

The broadband social acoustic signaling behavior of spinner and spotted dolphins.

Efforts to study the social acoustic signaling behavior of delphinids have traditionally been restricted to audio-range (<20 kHz) analyses. To explore the occurrence of communication signals at ultrasonic frequencies, broadband recordings of whistles and burst pulses were obtained from two commonly studied species of delphinids, the Hawaiian spinner dolphin (Stenella longirostris) and the Atlantic spotted dolphin (Stenella frontalis). Signals were quantitatively analyzed to establish their full bandwidth, to identify distinguishing characteristics between each species, and to determine how often they occur beyond the range of human hearing. Fundamental whistle contours were found to extend beyond 20 kHz only rarely among spotted dolphins, but with some regularity in spinner dolphins. Harmonics were present in the majority of whistles and varied considerably in their number, occurrence, and amplitude. Many whistles had harmonics that extended past 50 kHz and some reached as high as 100 kHz. The relative amplitude of harmonics and the high hearing sensitivity of dolphins to equivalent frequencies suggest that harmonics are biologically relevant spectral features. The burst pulses of both species were found to be predominantly ultrasonic, often with little or no energy below 20 kHz. The findings presented reveal that the social signals produced by spinner and spotted dolphins span the full range of their hearing sensitivity, are spectrally quite varied, and in the case of burst pulses are probably produced more frequently than reported by audio-range analyses.

Animals↗

Echo strength and density structure of Hawaiian mesopelagic boundary community patches.

A broadband sonar system and digital camera with strobe lights were mounted on a vertically profiling frame with a depth sensor. The echo strengths and densities of animals within individual mesopelagic boundary community patches were investigated as a function of depth. Time and distance from shore were also investigated. Simultaneous surface echosounder surveys permitted comparison of density estimates from two techniques. Echo strength values suggest nearshore boundary community animals are primarily myctophid fishes, which was confirmed by preliminary photographic evidence. Echo strength varied significantly as a function of distance from the shoreline and time. These measures of echo strength are important for estimating density from a surface echosounder. Density estimates from these revised echo strengths compare well with those made with echo highlight counting, which is independent of echo strength. These density measures suggest that previous density estimates were too low but do not change the conclusions of these studies. Vertical microstructure in density was apparent but animal size and compositional structure was not evident within a patch. Patch edges were abrupt, with no differences in the density or echo strength from patch interiors. These edges were generally straight, with a sharp drop in density to the background density of zero. Estimates of animal size as a function of time provide information about the diel migration patterns of these mesopelagic animals.

Journal Article↗

Acoustic backscattering by Hawaiian lutjanid snappers. 1. Target strength and swimbladder characteristics.

The target strengths and swimbladder morphology of six snapper species were investigated using broadband sonar, x rays, and swimbladder casts. Backscatter data were obtained using a frequency-modulated sweep (60-200 kHz) and a broadband, dolphinlike click (peak frequency 120 kHz) from live fish, mounted and rotated around each of their three axes. X rays revealed species-specific differences in the shape, size, and orientation of the swimbladders. The angle between the fish's dorsal aspect and the major axis of its swimbladder ranged from 3 degrees to 12 degrees and was consistent between individuals within a species. This angle had a one-to-one relationship with the angle at which the maximum dorsal aspect target strength was measured (r2 = 0.93), regardless of species. Maximum dorsal aspect target strength was correlated with length within species. However, the swimbladder modeled as an air-filled prolate spheroid with axes measured from the x rays of the swimbladder predicted maximum target strength significantly better than models based on fish length or swimbladder volume. For both the dorsal and lateral aspects, the prolate spheroid model's predictions were not significantly different from the measured target strengths (observed power >0.75) and were within 3 dB of the measured values. This model predicts the target strengths of all species equally well, unlike those based on length.

Acoustics↗