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Transformation of external-ear spectral cues into perceived delays by the big brown bat, Eptesicus fuscus.

The external-ear transfer function for big brown bats (Eptesicus fuscus) contains two prominent notches that vary from 30 to 55 kHz and from 70 to 100 kHz, respectively, as sound-source elevation moves from -40 to +10 degrees. These notches resemble a higher-frequency version of external-ear cues for vertical localization in humans and other mammals. However, they also resemble interference notches created in echoes when reflected sounds overlap at short time separations of 30-50 micros. Psychophysical experiments have shown that bats actually perceive small time separations from interference notches, and here we used the same technique to test whether external-ear notches are recognized as a corresponding time separation, too. The bats' performance reveals the elevation dependence of a time-separation estimate at 25-45 micros in perceived delay. Convergence of target-shape and external-ear cues onto echo spectra creates ambiguity about whether a particular notch relates to the object or to its location, which the bat could resolve by ignoring the presence of notches at external-ear frequencies. Instead, the bat registers the frequencies of notches caused by the external ear along with notches caused by the target's structure and employs spectrogram correlation and transformation (SCAT) to convert them all into a family of delay estimates that includes elevation.

Animals↗

Depth-dependent acoustic features of diving sperm whales (Physeter macrocephalus) in the Gulf of Mexico.

Three-dimensional dive trajectories of three sperm whales in the Gulf of Mexico have been obtained by measuring the relative arrival times and bearings of the animals' acoustic multipath reflections, using two elements of a towed hydrophone array deployed at an unknown depth and orientation. Within the first 6-12 min of the start of a dive, the intervals between successive "clicks" of all three whales corresponded closely with the two-way travel time of an acoustic pulse traveling vertically between the animals' position and the ocean bottom. The click spectra contained multiple peaks, including a faint band of energy originally centered near 10 kHz. As the animals descended over 500 m in depth, the center frequency of this band shifted to nearly 15 kHz, but subsequently remained near this value during the rest of the dive. This frequency shift is consistent with that expected from energy scattering from an ensemble of incompressible small-scale air-filled resonators, with diameters on the order of 4 mm. One possible candidate for such an ensemble is proposed to reside in the collapsed frontal sac of the animal. A comparison of the received levels for the bottom and direct multipath arrivals indicates that the whales' acoustic directivity must range between 10-30 dB in the 5-20-kHz region.

Acoustics↗

The effect of a low-frequency sound source (acoustic thermometry of the ocean climate) on the diving behavior of juvenile northern elephant seals, Mirounga angustirostris.

Changes in the diving behavior of individual free-ranging juvenile northern elephant seals, Mirounga angustirostris, exposed to the acoustic thermometry of the ocean climate (ATOC) sound source were examined using data loggers. Data loggers were attached to the animals and measured swim speed, maximum depth of dive, dive duration, surface interval, descent and ascent rate, and descent and ascent angle along with sound pressure level (SPL). The ATOC sound source was at a depth of 939 m and transmitted at 195 dB re: 1 microPa at 1 m centered at 75 Hz with a 37.5-Hz bandwidth. Sound pressure levels (SPL) measured at the seal during transmissions averaged 128 dB and ranged from 118 to 137 dB re: 1 microPa for the 60-90 Hz band, in comparison to ambient levels of 87-107 dB within this band. In no case did an animal end its dive or show any other obvious change in behavior upon exposure to the ATOC sound. Subtle changes in diving behavior were detected, however. During exposure, deviations in descent rate were greater than 1 s.d. of the control mean in 9 of 14 seals. Dive depth increased and descent velocity increased in three animals, ascent velocity decreased in two animals, ascent rate increased in one animal and decreased in another, and dive duration decreased in only one animal. There was a highly significant positive correlation between SPL and descent rate. The biological significance of these subtle changes is likely to be minimal. This is the first study to quantify behavioral responses of an animal underwater with simultaneous measurements of SPL of anthropogenic sounds recorded at the animal.

Acoustics↗

Time-frequency model for echo-delay resolution in wideband biosonar.

A time/frequency model of the bat's auditory system was developed to examine the basis for the fine (approximately 2 micros) echo-delay resolution of big brown bats (Eptesicus fuscus), and its performance at resolving closely spaced FM sonar echoes in the bat's 20-100-kHz band at different signal-to-noise ratios was computed. The model uses parallel bandpass filters spaced over this band to generate envelopes that individually can have much lower bandwidth than the bat's ultrasonic sonar sounds and still achieve fine delay resolution. Because fine delay separations are inside the integration time of the model's filters (approximately 250-300 micros), resolving them means using interference patterns along the frequency dimension (spectral peaks and notches). The low bandwidth content of the filter outputs is suitable for relay of information to higher auditory areas that have intrinsically poor temporal response properties. If implemented in fully parallel analog-digital hardware, the model is computationally extremely efficient and would improve resolution in military and industrial sonar receivers.

Animals↗

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↗

Outer hair cell piezoelectricity: frequency response enhancement and resonance behavior.

Stretching or compressing an outer hair cell alters its membrane potential and, conversely, changing the electrical potential alters its length. This bi-directional energy conversion takes place in the cell's lateral wall and resembles the direct and converse piezoelectric effects both qualitatively and quantitatively. A piezoelectric model of the lateral wall has been developed that is based on the electrical and material parameters of the lateral wall. An equivalent circuit for the outer hair cell that includes piezoelectricity shows a greater admittance at high frequencies than one containing only membrane resistance and capacitance. The model also predicts resonance at ultrasonic frequencies that is inversely proportional to cell length. These features suggest all mammals use outer hair cell piezoelectricity to support the high-frequency receptor potentials that drive electromotility. It is also possible that members of some mammalian orders use outer hair cell piezoelectric resonance in detecting species-specific vocalizations.

Animals↗

Evaluation of an auditory model for echo delay accuracy in wideband biosonar.

In a psychophysical task with echoes that jitter in delay, big brown bats can detect changes as small as 10-20 ns at an echo signal-to-noise ratio of approximately 49 dB and 40 ns at approximately 36 dB. This performance is possible to achieve with ideal coherent processing of the wideband echoes, but it is widely assumed that the bat's peripheral auditory system is incapable of encoding signal waveforms to represent delay with the requisite precision or phase at ultrasonic frequencies. This assumption was examined by modeling inner-ear transduction with a bank of parallel bandpass filters followed by low-pass smoothing. Several versions of the filterbank model were tested to learn how the smoothing filters, which are the most critical parameter for controlling the coherence of the representation, affect replication of the bat's performance. When tested at a signal-to-noise ratio of 36 dB, the model achieved a delay acuity of 83 ns using a second-order smoothing filter with a cutoff frequency of 8 kHz. The same model achieved a delay acuity of 17 ns when tested with a signal-to-noise ratio of 50 dB. Jitter detection thresholds were an order of magnitude worse than the bat for fifth-order smoothing or for lower cutoff frequencies. Most surprising is that effectively coherent reception is possible with filter cutoff frequencies well below any of the ultrasonic frequencies contained in the bat's sonar sounds. The results suggest that only a modest rise in the frequency response of smoothing in the bat's inner ear can confer full phase sensitivity on subsequent processing and account for the bat's fine acuity or delay.

Animals↗

Click sounds produced by cod (Gadus morhua).

Conspicuous sonic click sounds were recorded in the presence of cod (Gadus morhua), together with either harp seals (Pagophilus groenlandicus), hooded seals (Cystophora cristata) or a human diver in a pool. Similar sounds were never recorded in the presence of salmon (Salmo salar) together with either seal species, or from either seal or fish species when kept separately in the pool. It is concluded that cod was the source of these sounds and that the clicks were produced only when cod were approached by a swimming predatorlike body. The analyzed click sounds (n = 377) had the following characteristics (overall averages +/- S.D.): peak frequency = 5.95 +/- 2.22 kHz; peak-to-peak duration = 0.70 +/- 0.45 ms; sound pressure level (received level) = 153.2 +/- 7.0 dB re 1 microPa at 1 m. At present the mechanism and purpose of these clicks is not known. However, the circumstances under which they were recorded and some observations on the behavior of the seals both suggest that the clicks could have a predator startling function.

Animal Communication↗

Localization of aerial broadband noise by pinnipeds.

Although many pinnipeds (seals, sea lions, and walruses) emit broadband calls on land as part of their communication system, few studies have addressed these animals' ability to localize aerial broadband sounds. In this study, the aerial sound localization acuities of a female northern elephant seal (Mirounga angustirostris), a male harbor seal (Phoca vitulina), and a female California sea lion (Zalophus californianus) were measured in the horizontal plane. The stimulus was broadband white noise that was band pass filtered between 1.2 and 15 kHz. Testing was conducted in a hemi-anechoic chamber using a left/right forced choice procedure to measure the minimum audible angle (MAA) for each subject. MAAs were defined as half the angular separation of two sound sources bisected by a subject's midline that corresponded to 75% correct discrimination. MAAs were 4.7 degrees, 3.6 degrees, and 4.2 degrees for the northern elephant seal, harbor seal, and California sea lion, respectively. These results demonstrate that individuals of these pinniped species have sound localization abilities comparable to the domestic cat and rhesus macaque. The acuity differences between our subjects were small and not predicted by head size. These results likely reflect the relatively acute general abilities of pinnipeds to localize aerial broadband signals.

Animals↗

Estimated transmission beam pattern of clicks recorded from free-ranging white-beaked dolphins (Lagenorhynchus albirostris).

Recordings were made from white-beaked dolphins in Icelandic waters using a four-hydrophone array in a star configuration. The acoustic signals were amplified and sampled to a hard disk at a rate of 800 kHz per channel. The 3 and 10 dB beamwidths were calculated to be 8 degrees and 10 degrees, respectively, indicating a narrower transmission beam for white-beaked dolphins than that reported for bottlenose dolphins (Tursiops truncatus). The beamwidth was more similar to that found for belugas (Delphinapterus lucas). The measured beam pattern included large side lobes, perhaps due to the inclusion of off-axis clicks, even after applying several criteria to select only on-axis clicks. The directivity index was calculated to be 18 dB when using all data for angles from 0 degrees-50 degrees. The calculated sound radiation from a circular piston with a radius of 6 cm driven by a white-beaked dolphin click had a beam pattern very similar to the measured beam pattern for the main transmission lobe of the white-beaked dolphin. The directivity index was 29 dB. This is the first attempt to estimate the directionality index of dolphins in the field.

Acoustic Stimulation↗

Neuro-computational processing of moving sonar echoes classifies and localizes foliage.

Echoes from in situ tree trunks, similar to those observed by flying bats, are processed. A moving sonar converts echoes into spike sequences and applies neural-computational methods to classify objects and estimate passing range. Two classes of tree trunks act as retro-reflectors that generate strong echoes (SEs), identified by a locally dense spike pattern. Linear drive-by sonar trajectories cause SEs to follow hyperbolic curves specified by passing range. A glint is a collection of consecutive range readings matching expected values on a specific hyperbolic curve. Passing-range detectors compare successive SE data with expected values in a table and tally coincidences. Counters increment when coincidences occur and decrement when they do not. A glint terminates after tallying a sufficient number of coincidences and coincidence failure occurs in the maximum-count detector. Reflector roughness, deviations in sonar trajectory, and echo jitter necessitate a coincidence window to define matches. Short windows identify small glints over piecewise linear sonar trajectories, while long windows accommodate deviations in sonar speed and trajectory, and associate multiple glints observed with shorter windows. The minimum coincidence window size yielding glints classify smooth and rough retro-reflectors.

Acoustic Stimulation↗

Geometry of locating sounds from differences in travel time: isodiachrons.

Calling animals may be located from measurements of the differences in acoustic travel time at pairs of receivers. For inhomogeneous fields of speed, locations can be made with better accuracy when the location algorithm allows the speed to vary from path to path. A new geometrical shape, called an isodiachron, is described. It is the locus of points corresponding to a constant difference in travel time along straight paths between the animal and two receivers. Its properties allow an interpretation for locations when the speed differs from path to path. An algorithm has been developed for finding the location of calling animals by intersecting isodiachrons from data collected at pairs of receivers. When the sound speed field is spatially homogeneous, isodiachrons become hyperboloids. Unlike a hyperboloid that extends to infinity, an isodiachron is confined to a finite region of space when the speeds differ between the animal and each of two receivers. Its shape is significantly different than a hyperboloid for cases of practical interest. Isodiachrons can be used to better understand locations of calling animals and other sounds in the sea, Earth, and air.

Air↗

A numerical study of the role of the tragus in the big brown bat.

A comprehensive characterization of the spatial sensitivity of an outer ear from a big brown bat (Eptesicus fuscus) has been obtained using numerical methods and visualization techniques. Pinna shape information was acquired through x-ray microtomography. It was used to set up a finite-element model of diffraction from which directivities were predicted by virtue of forward wave-field projections based on a Kirchhoff integral formulation. Digital shape manipulation was used to study the role of the tragus in detailed numerical experiments. The relative position between tragus and pinna aperture was found to control the strength of an extensive asymmetric sidelobe which points in a frequency-dependent direction. An upright tragus position resulted in the strongest sidelobe sensitivity. Using a bootstrap validation paradigm, the results were found to be robust against small perturbations of the finite-element mesh boundaries. Furthermore, it was established that a major aspect of the tragus effect (position dependence) can be studied in a simple shape model, an obliquely truncated horn augmented by a flap representing the tragus. In the simulated wave field around the outer-ear structure, strong correlates of the tragus rotation were identified, which provide a direct link to the underlying physical mechanism.

Animals↗

Three-dimensional beam pattern of regular sperm whale clicks confirms bent-horn hypothesis.

The three-dimensional beam pattern of a sperm whale (Physeter macrocephalus) tagged in the Ligurian Sea was derived using data on regular clicks from the tag and from hydrophones towed behind a ship circling the tagged whale. The tag defined the orientation of the whale, while sightings and beamformer data were used to locate the whale with respect to the ship. The existence of a narrow, forward-directed P1 beam with source levels exceeding 210 dBpeak re: 1 microPa at 1 m is confirmed. A modeled forward-beam pattern, that matches clicks >20 degrees off-axis, predicts a directivity index of 26.7 dB and source levels of up to 229 dBpeak re: 1 microPa at 1 m. A broader backward-directed beam is produced by the P0 pulse with source levels near 200 dBpeak re: 1 microPa at 1 m and a directivity index of 7.4 dB. A low-frequency component with source levels near 190 dBpeak re: 1 microPa at 1 m is generated at the onset of the P0 pulse by air resonance. The results support the bent-horn model of sound production in sperm whales. While the sperm whale nose appears primarily adapted to produce an intense forward-directed sonar signal, less-directional click components convey information to conspecifics, and give rise to echoes from the seafloor and the surface, which may be useful for orientation during dives.

Acoustics↗

Are high perches in the blackcap Sylvia atricapilla song or listening posts? A sound transmission study.

Birds often sing from high perches referred to as song posts. However, birds also listen and keep a lookout from these perches. We used a sound transmission experiment to investigate the changes for receiving and sending conditions that a territorial songbird may experience by moving upwards in the vegetation. Representative song elements of the blackcap Sylvia atricapilla were transmitted in a forest habitat in spring using a complete factorial design with natural transmission distances and speaker and microphone heights. Four aspects of sound degradation were quantified: signal-to-noise ratio, excess attenuation, distortion within the sounds determined as a blur ratio, and prolongation of the sounds with "tails" of echoes determined as a tail-to-signal ratio. All four measures indicated that degradation decreased with speaker and microphone height. However, the decrease was considerably higher for the microphone than for the speaker. This suggests that choosing high perches in a forest at spring results in more benefits to blackcaps in terms of improved communication conditions when they act as receivers than as senders.

Amplifiers, Electronic↗