The transmission beam pattern of an echolocating harbor porpoise (Phocoena phocoena) was measured in both the vertical and horizontal planes. An array of seven Brüel and Kjaer 8103 hydrophones connected to an amplifier-line driver module was used to measure the beam patterns. The porpoise was trained to station in a hoop and echolocate a cylindrical target located at a range between 7 and 9 m while the array was located 2 m in front of the hoop. The 3-dB beamwidth in both the vertical and horizontal planes was the same at approximately 16 degrees and the beam was pointed toward the forward direction. The individual hydrophones in both the vertical and horizontal arrays measured signal waveforms that were similar throughout the 40-degree span of the array. The porpoise emitted signals with intervals that were 20 to 35 ms longer than the round trip travel time between the animal and the target. The average source level, peak frequency, and bandwidth were 157 dB, 128 kHz, and 16 kHz, respectively.
The underwater hearing sensitivity of a two-year-old harbor porpoise 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 narrow-band frequency-modulated signals having center frequencies between 250 Hz and 180 kHz. The resulting audiogram was U-shaped with the range of best hearing (defined as 10 dB within maximum sensitivity) from 16 to 140 kHz, with a reduced sensitivity around 64 kHz. Maximum sensitivity (about 33 dB re 1 microPa) occurred between 100 and 140 kHz. This maximum sensitivity range corresponds with the peak frequency of echolocation pulses produced by harbor porpoises (120-130 kHz). Sensitivity falls about 10 dB per octave below 16 kHz and falls off sharply above 140 kHz (260 dB per octave). Compared to a previous audiogram of this species (Andersen, 1970), the present audiogram shows less sensitive hearing between 2 and 8 kHz and more sensitive hearing between 16 and 180 kHz. This harbor porpoise has the highest upper-frequency limit of all odontocetes investigated. The time it took for the porpoise to move its head 22 cm after the signal onset (movement time) was also measured. It increased from about 1 s at 10 dB above threshold, to about 1.5 s at threshold.
The underwater hearing sensitivity of a young male harbor porpoise for tonal signals of various signal durations was quantified by using a behavioral psychophysical technique. The animal was trained to respond only when it detected an acoustic signal. Fifty percent detection thresholds were obtained for tonal signals (15 frequencies between 0.25-160 kHz, durations 0.5-5000 ms depending on the frequency; 134 frequency-duration combinations in total). Detection thresholds were quantified by varying signal amplitude by the 1-up 1-down staircase method. The hearing thresholds increased when the signal duration fell below the time constant of integration. The time constants, derived from an exponential model of integration [Plomp and Bouman, J. Acoust. Soc. Am. 31, 749-758 (1959)], varied from 629 ms at 2 kHz to 39 ms at 64 kHz. The integration times of the porpoises were similar to those of other mammals including humans, even though the porpoise is a marine mammal and a hearing specialist. The results enable more accurate estimations of the distances at which porpoises can detect short-duration environmental tonal signals. The audiogram thresholds presented by Kastelein et al. [J. Acoust. Soc. Am. 112, 334-344 (2002)], after correction for the frequency bandwidth of the FM signals, are similar to the results of the present study for signals of 1500 ms duration. Harbor porpoise hearing is more sensitive between 2 and 10 kHz, and less sensitive above 10 kHz, than formerly believed.
Safety criteria for underwater sound produced during offshore pile driving are needed to protect marine mammals. A harbor porpoise was exposed to fatiguing noise at 18 sound pressure level (SPL) and duration combinations. Its temporary hearing threshold shift (TTS) and hearing recovery were quantified with a psychoacoustic technique. Octave-band white noise centered at 4 kHz was the fatiguing stimulus at three mean received SPLs (124, 136, and 148 dB re 1 μPa) and at six durations (7.5, 15, 30, 60, 120, and 240 min). Approximate received sound exposure levels (SELs) varied between 151 and 190 dB re 1 μPa(2) s. Hearing thresholds were determined for a narrow-band frequency-swept sine wave (3.9-4.1 kHz; 1 s) before exposure to the fatiguing noise, and at 1-4, 4-8, 8-12, 48, and 96 min after exposure. The lowest SEL (151 dB re 1 μPa(2) s) which caused a significant TTS(1-4) was due to exposure to an SPL of 124 dB re 1 μPa for 7.5 min. The maximum TTS(1-4), induced after a 240 min exposure to 148 dB re 1 μPa, was around 15 dB at a SEL of 190 dB re 1 μPa(2) s. Recovery time following TTS varied between 4 min and under 96 min, depending on the exposure level, duration, and the TTS induced.
SUMMARY 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°. 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.
The underwater hearing sensitivities of two 1-year-old female harbor seals were quantified in a pool built for acoustic research, using a behavioral psychoacoustic technique. The animals were trained to respond when they detected an acoustic signal and not to respond when they did not (go/no-go response). Pure tones (0.125-0.25 kHz) and narrowband frequency modulated (tonal) signals (center frequencies 0.5-100 kHz) of 900 ms duration were tested. Thresholds at each frequency were measured using the up-down staircase method and defined as the stimulus level resulting in a 50% detection rate. The audiograms of the two seals did not differ statistically: both plots showed the typical mammalian U-shape, but with a wide and flat bottom. Maximum sensitivity (54 dB re 1 microPa, rms) occurred at 1 kHz. The frequency range of best hearing (within 10 dB of maximum sensitivity) was from 0.5 to 40 kHz (6(1/3) octaves). Higher hearing thresholds (indicating poorer sensitivity) were observed below 1 and above 40 kHz. Thresholds below 4 kHz were lower than those previously described for harbor seals, which demonstrates the importance of using quiet facilities, built specifically for acoustic research, for hearing studies in marine mammals. The results suggest that under unmasked conditions many anthropogenic noise sources and sounds from conspecifics are audible to harbor seals at greater ranges than formerly believed.
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