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.
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