Threshold received sound pressure levels of single 1-2 kHz and 6-7 kHz up-sweeps and down-sweeps causing startle responses in a harbor porpoise (Phocoena phocoena)

Mid-frequency and low-frequency sonar systems produce frequency-modulated sweeps which may affect harbor porpoises. To study the effect of sweeps on behavioral responses (specifically "startle" responses, which we define as sudden changes in swimming speed and/or direction), a harbor porpoise in a large pool was exposed to three pairs of sweeps: a 1-2 kHz up-sweep was compared with a 2-1 kHz down-sweep, both with and without harmonics, and a 6-7 kHz up-sweep was compared with a 7-6 kHz down-sweep without harmonics. Sweeps were presented at five spatially averaged received levels (mRLs; 6 dB steps; identical for the up-sweep and down-sweep of each pair). During sweep presentation, startle responses were recorded. There was no difference in the mRLs causing startle responses for up-sweeps and down-sweeps within frequency pairs. For 1-2 kHz sweeps without harmonics, a 50% startle response rate occurred at mRLs of 133 dB re 1 μPa; for 1-2 kHz sweeps with strong harmonics at 99 dB re 1 μPa; for 6-7 kHz sweeps without harmonics at 101 dB re 1 μPa. Low-frequency (1-2 kHz) active naval sonar systems without harmonics can therefore operate at higher source levels than mid-frequency (6-7 kHz) active sonar systems without harmonics, with similar startle effects on porpoises.     

Hearing thresholds of a harbor porpoise (Phocoena phocoena) for helicopter dipping sonar signals (1.43-1.33 kHz) (L)

Helicopter long range active sonar (HELRAS), a "dipping" sonar system used by lowering transducer and receiver arrays into water from helicopters, produces signals within the functional hearing range of many marine animals, including the harbor porpoise. The distance at which the signals can be heard is unknown, and depends, among other factors, on the hearing sensitivity of the species to these particular signals. Therefore, the hearing thresholds of a harbor porpoise for HELRAS signals were quantified by means of a psychophysical technique. Detection thresholds were obtained for five 1.25 s simulated HELRAS signals, varying in their harmonic content and amplitude envelopes. The 50% hearing thresholds for the different signals were similar: 76 dB re 1 μPa (broadband sound pressure level, averaged over the signal duration). The detection thresholds were similar to those found in the same porpoise for tonal signals in the 1-2 kHz range measured in a previous study. Harmonic distortion, which occurred in three of the five signals, had little influence on their audibility. The results of this study, combined with information on the source level of the signal, the propagation conditions and ambient noise levels, allow the calculation of accurate estimates of the distances at which porpoises can detect HELRAS signals.     

Effect of broadband-noise masking on the behavioral response of a harbor porpoise (Phocoena phocoena) to 1-s duration 6-7 kHz sonar up-sweeps

Naval sonar systems produce signals which may affect the behavior of harbor porpoises, though their effect may be reduced by ambient noise. To show how natural ambient noise influences the effect of sonar sweeps on porpoises, a porpoise in a pool was exposed to 1-s duration up-sweeps, similar in frequency range (6-7 kHz) to those of existing naval sonar systems. The sweep signals had randomly generated sweep intervals of 3-7 s (duty cycle: 19%). Behavioral parameters during exposure to signals were compared to those during baseline periods. The sessions were conducted under five background noise conditions: the local normal ambient noise and four conditions mimicking the spectra for wind-generated noise at Sea States 2-8. In all conditions, the sweeps caused the porpoise to swim further away from the transducer, surface more often, swim faster, and breathe more forcefully than during the baseline periods. However, the higher the background noise level, the smaller the effects of the sweeps on the surfacing behavior of the porpoise. Therefore, the effects of naval sonar systems on harbor porpoises are determined not only by the received level of the signals and the hearing sensitivity of the animals but also by the background noise.     

Audiogram of a harbor porpoise (Phocoena phocoena) measured with narrow-band frequency-modulated signals

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 effect of signal duration on the underwater detection thresholds of a harbor porpoise (Phocoena phocoena) for single frequency-modulated tonal signals between 0.25 and 160 kHz

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.     

Noise-induced temporary threshold shift and recovery in Yangtze finless porpoises Neophocaena phocaenoides asiaeorientalis

In Yangtze finless porpoises Neophocaena phocaenoides asiaeorientalis, the effects of fatiguing noise on hearing thresholds at frequencies of 32, 45, 64, and 128 kHz were investigated. The noise parameters were: 0.5-oct bandwidth, -1 to +0.5 oct relative to the test frequency, 150 dB re 1 μPa (140-160 dB re 1 μPa in one measurement series), with 1-30 min exposure time. Thresholds were evaluated using the evoked-potential technique allowing the tracing of threshold variations with a temporal resolution better than 1 min. The most effective fatiguing noise was centered at 0.5 octave below the test frequency. The temporary threshold shift (TTS) depended on the frequencies of the fatiguing noise and test signal: The lower the frequencies, the bigger the noise effect. The time-to-level trade of the noise effect was incomplete: the change of noise level by 20 dB resulted in a change of TTS level by nearly 20 dB, whereas the tenfold change of noise duration resulted in a TTS increase by 3.8-5.8 dB.     

圈养瓶鼻海豚对20 kHz连续声信号的行为响应研究

将20 kHz连续声信号作为刺激信号,测试了厦门某海湾圈养的两只瓶鼻海豚对该信号的行为变化。通过对比信号发射期与间歇期海豚相对声源的水面距离、露出水面的次数以及水下发出的click定位声信号的数目等变化,判断发射信号对海豚行为的影响。给出了瓶鼻海豚对测试信号产生躲避行为的声压级门限(154±2 dB re 1μPa,rms),并与鼠海豚的躲避声压门限级进行了对比。结果表明:信号发射期,瓶鼻海豚移离了声源位置,增加了露出水面的次数,水下发出click声信号的次数明显减少。因此,瓶鼻海豚对20kHz连续信号产生了行为改变。      

Behavior of captive herring exposed to naval sonar transmissions (1.0-1.6 kHz) throughout a yearly cycle

Atlantic herring, Clupea harengus, is a hearing specialist, and several studies have demonstrated strong responses to man-made noise, for example, from an approaching vessel. To avoid negative impacts from naval sonar operations, a set of studies of reaction patters of herring to low-frequency (1.0-1.5 kHz) naval sonar signals has been undertaken. This paper presents herring reactions to sonar signals and other stimuli when kept in captivity under detailed acoustic and video monitoring. Throughout the experiment, spanning three seasons of a year, the fish did not react significantly to sonar signals from a passing frigate, at received root-mean-square sound-pressure level (SPL) up to 168 dB re 1 μPa. In contrast, the fish did exhibit a significant diving reaction when exposed to other sounds, with a much lower SPL, e.g., from a two-stroke engine. This shows that the experimental setup is sensitive to herring reactions when occurring. The lack of herring reaction to sonar signals is consistent with earlier in situ behavioral studies. The complexity of the behavioral reactions in captivity underline the need for better understanding of the causal relationship between stimuli and reaction patterns of fish.     

The hearing threshold of a harbor porpoise (Phocoena phocoena) for impulsive sounds (L)

The distance at which harbor porpoises can hear underwater detonation sounds is unknown, but depends, among other factors, on the hearing threshold of the species for impulsive sounds. Therefore, the underwater hearing threshold of a young harbor porpoise for an impulsive sound, designed to mimic a detonation pulse, was quantified by using a psychophysical technique. The synthetic exponential pulse with a 5?ms time constant was produced and transmitted by an underwater projector in a pool. The resulting underwater sound, though modified by the response of the projection system and by the pool, exhibited the characteristic features of detonation sounds: A zero to peak sound pressure level of at least 30?dB (re 1?s(-1)) higher than the sound exposure level, and a short duration (34?ms). The animal's 50% detection threshold for this impulsive sound occurred at a received unweighted broadband sound exposure level of 60?dB re 1?μPa(2)s. It is shown that the porpoise's audiogram for short-duration tonal signals [Kastelein et al., J. Acoust. Soc. Am. 128, 3211-3222 (2010)] can be used to estimate its hearing threshold for impulsive sounds.     

Near-threshold equal-loudness contours for harbor seals (Phoca vitulina) derived from reaction times during underwater audiometry: a preliminary study

Equal-loudness functions describe relationships between the frequencies of sounds and their perceived loudness. This pilot study investigated the possibility of deriving equal-loudness contours based on the assumption that sounds of equal perceived loudness elicit equal reaction times (RTs). During a psychoacoustic underwater hearing study, the responses of two young female harbor seals to tonal signals between 0.125 and 100 kHz were filmed. Frame-by-frame analysis was used to quantify RT (the time between the onset of the sound stimulus and the onset of movement of the seal away from the listening station). Near-threshold equal-latency contours, as surrogates for equal-loudness contours, were estimated from RT-level functions fitted to mean RT data. The closer the received sound pressure level was to the 50% detection hearing threshold, the more slowly the animals reacted to the signal (RT range: 188-982 ms). Equal-latency contours were calculated relative to the RTs shown by each seal at sound levels of 0, 10, and 20 dB above the detection threshold at 1 kHz. Fifty percent detection thresholds are obtained with well-trained subjects actively listening for faint familiar sounds. When calculating audibility ranges of sounds for harbor seals in nature, it may be appropriate to consider levels 20 dB above this threshold.     

Behavioral avoidance threshold level of a harbor porpoise (Phocoena phocoena) for a continuous 50 kHz pure tone

The use of ultrasonic sounds in alarms for gillnets may be advantageous, but the deterring effects of ultrasound on porpoises are not well understood. Therefore a harbor porpoise in a large floating pen was subjected to a continuous 50 kHz pure tone with a source level of 122+/-3 dB (re 1 microPa, rms). When the test signal was switched on during test periods, the animal moved away from the sound source. Its respiration rate was similar to that during baseline periods, when the sound was switched off. The behavior of the porpoise was related to the sound pressure level distribution in the pen. The sound level at the animal's average swimming location during the test periods was approximately 107+/-3 dB (re 1 microPa, rms). The avoidance threshold sound pressure level for a continuous 50 kHz pure tone for this porpoise, in the context of this study, is estimated to be 108+/-3 dB (re 1 microPa, rms). This study demonstrates that porpoises may be deterred from an area by high frequency sounds that are not typically audible to fish and pinnipeds and would be less likely masked by ambient noise.     

Behavioral responses of two captive bottlenose dolphins (Tursiops truncatus) to a continuous 50 kHz tone

At present, the fundamental frequencies of signals of most commercially available acoustic alarms to deter small cetaceans are below 20 kHz, but it is not well ascertained whether higher frequencies have a deterrent effect on bottlenose dolphins (Tursiops truncatus). Two captive bottlenose dolphins housed in a floating pen were subjected to a continuous pure tone at 50 kHz with a source level of 160 ± 2 dB (re 1 μPa, rms). The behavioral responses of dolphins were judged by comparing surfacing distance relative to the sound source, number of surfacings, and number of echolocation clicks produced, during forty 15 min baseline periods with forty 15 min test periods (four sessions per day, 40 sessions in total). On all 10 study days, surfacing distance and the number of surfacings increased while click production decreased during broadcasts of test sound. The avoidance threshold sound pressure level for a continuous 50 kHz tone for the bottlenose dolphins, in the context of this study, was estimated to be 144 ± 2 dB (re 1 μPa, rms). The results indicated that a continuous 50 kHz tonal signal can deter bottlenose dolphins from an area.     

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.     

Temporary threshold shift in bottlenose dolphins (Tursiops truncatus) exposed to mid-frequency tones

A behavioral response paradigm was used to measure hearing thresholds in bottlenose dolphins before and after exposure to 3 kHz tones with sound exposure levels (SELs) from 100 to 203 dB re 1 microPa2 s. Experiments were conducted in a relatively quiet pool with ambient noise levels below 55 dB re 1 microPa2/Hz at frequencies above 1 kHz. Experiments 1 and 2 featured 1-s exposures with hearing tested at 4.5 and 3 kHz, respectively. Experiment 3 featured 2-, 4-, and 8-s exposures with hearing tested at 4.5 kHz. For experiment 2, there were no significant differences between control and exposure sessions. For experiments 1 and 3, exposures with SEL=197 dB re 1 microPa2 s and SEL > or = 195 dB re 1 microPa2 s, respectively, resulted in significantly higher TTS4 than control sessions. For experiment 3 at SEL= 195 dB re 1 microPa2 s, the mean TTS4 was 2.8 dB. These data are consistent with prior studies of TTS in dolphins exposed to pure tones and octave band noise and suggest that a SEL of 195 dB re 1 microPa2 s is a reasonable threshold for the onset of TTS in dolphins and white whales exposed to midfrequency tones.     

The influence of signal parameters on the sound source localization ability of a harbor porpoise (Phocoena phocoena)

It is unclear how well harbor porpoises can locate sound sources, and thus can locate acoustic alarms on gillnets. Therefore the ability of a porpoise to determine the location of a sound source was determined. The animal was trained to indicate the active one of 16 transducers in a 16-m-diam circle around a central listening station. The duration and received level of the narrowband frequency-modulated signals (center frequencies 16, 64 and 100 kHz) were varied. The animal's localization performance increased when the signal duration increased from 600 to 1000 ms. The lower the received sound pressure level (SPL) of the signal, the harder the animal found it to localize the sound source. When pulse duration was long enough (approximately 1 s) and the received SPLs of the sounds were high (34-50 dB above basic hearing thresholds or 3-15 dB above the theoretical masked detection threshold in the ambient noise condition of the present study), the animal could locate sounds of the three frequencies almost equally well. The porpoise was able to locate sound sources up to 124 degrees to its left or right more easily than sounds from behind it.     

Effects of mid-frequency active sonar on hearing in fish

Caged fish were exposed to sound from mid-frequency active (MFA) transducers in a 5 × 5 planar array which simulated MFA sounds at received sound pressure levels of 210 dB SPL(re 1 μPa). The exposure sound consisted of a 2 s frequency sweep from 2.8 to 3.8 kHz followed by a 1 s tone at 3.3 kHz. The sound sequence was repeated every 25 s for five repetitions resulting in a cumulative sound exposure level (SEL(cum)) of 220 dB re 1 μPa(2) s. The cumulative exposure level did not affect the hearing sensitivity of rainbow trout, a species whose hearing range is lower than the frequencies in the presented MFA sound. In contrast, one cohort of channel catfish showed a statistically significant temporary threshold shift of 4-6 dB at 2300 Hz, but not at lower tested frequencies, whereas a second cohort showed no change. It is likely that this threshold shift resulted from the frequency spectrum of the MFA sound overlapping with the upper end of the hearing frequency range of the channel catfish. The observed threshold shifts in channel catfish recovered within 24 h. There was no mortality associated with the MFA sound exposure used in this test.     

Receiving beam patterns in the horizontal plane of a harbor porpoise (Phocoena phocoena)

Receiving beam patterns of a harbor porpoise were measured in the horizontal plane, using narrow-band frequency modulated signals with center frequencies of 16, 64, and 100 kHz. Total signal duration was 1000 ms, including a 200 ms rise time and 300 ms fall time. The harbor porpoise was trained to participate in a psychophysical test and stationed itself horizontally in a specific direction in the center of a 16-m-diameter circle consisting of 16 equally-spaced underwater transducers. The animal's head and the transducers were in the same horizontal plane, 1.5 m below the water surface. The go/no-go response paradigm was used; the animal left the listening station when it heard a sound signal. The method of constants was applied. For each transducer the 50% detection threshold amplitude was determined in 16 trials per amplitude, for each of the three frequencies. The beam patterns were not symmetrical with respect to the midline of the animal's body, but had a deflection of 3-7 degrees to the right. The receiving beam pattern narrowed with increasing frequency. Assuming that the pattern is rotation-symmetrical according to an average of the horizontal beam pattern halves, the receiving directivity indices are 4.3 at 16 kHz, 6.0 at 64 kHz, and 11.7 dB at 100 kHz. The receiving directivity indices of the porpoise were lower than those measured for bottlenose dolphins. This means that harbor porpoises have wider receiving beam patterns than bottlenose dolphins for the same frequencies. Directivity of hearing improves the signal-to-noise ratio and thus is a tool for a better detection of certain signals in a given ambient noise condition.     

Comparison between visual and passive acoustic detection of finless porpoises in the Yangtze River, China

Recently, sonar signals and other sounds produced by cetaceans have been used for acoustic detection of individuals and groups in the wild. However, the detection probability ascertained by concomitant visual survey has not been demonstrated extensively. The finless porpoises (Neophocaena phocaenoides) have narrow band and high-frequency sonar signals, which are distinctive from background noises. Underwater sound monitoring with hydrophones (B&K8103) placed along the sides of a research vessel, concurrent with visual observations was conducted in the Yangtze River from Wuhan to Poyang Lake in 1998 in China. The peak to peak detection threshold was set at 133 dB re 1 ,EPa. With this threshold level, porpoises could be detected reliably within 300 m of the hydrophone. In a total of 774-km cruise, 588 finless porpoises were sighted by visual observation and 44 ,864 ultrasonic pulses were recorded by the acoustical observation system. The acoustic monitoring system could detect the presence of the finless porpoises 82% of the time. A false alarm in the system occurred with a frequency of 0.9%. The high-frequency acoustical observation is suggested as an effective method for field surveys of small cetaceans, which produce high-frequency sonar signals.     

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.     

Localization and source level estimates of black drum (Pogonias cromis) calls

A four hydrophone linear array was used to localize calling black drum and estimate source levels and signal propagation. A total of 1025 source level estimates averaged 165 dB(RMS) relative (re:) 1 μPa (standard deviation (SD)=1.0). The authors suggest that the diverticulated morphology of the black drum swimbladder increase the bladder's surface area, thus contributing to sound amplitude. Call energy was greatest in the fundamental frequency (94 Hz) followed by the second (188 Hz) and third harmonics (282 Hz). A square root model best described propagation of the entire call, and separately the fundamental frequency and second harmonic. A logarithmic model best described propagation of the third harmonic which was the only component to satisfy the cut-off frequency equation. Peak auditory sensitivity was 300 Hz at a 94 dB re: 1 μPa threshold based on auditory evoked potential measurements of a single black drum. Based on mean RMS source level, signal propagation, background levels, and hearing sensitivity, the communication range of black drum was estimated at 33-108 m and was limited by background levels not auditory sensitivity. This estimate assumed the source and receiver were at approximately 0.5 m above the bottom. Consecutive calls of an individual fish localized over 59 min demonstrated a mean calling period of 3.6 s (SD=0.48), mean swimming speed of 0.5 body lengths/s, and a total distance swam of 1035 m.