Detection and discrimination of simulated motion of auditory targets in the horizontal plane

Three experiments investigated subjects' ability to detect and discriminate the simulated horizontal motion of auditory targets in an anechoic environment. "Moving" stimuli were produced by dynamic application of stereophonic balancing algorithms to a two-loudspeaker system with a 30 degree separation. All stimuli were 500-Hz tones. In experiment 1, subjects had to discriminate a left-to-right moving stimulus from a stationary stimulus pulsed for the same duration (300 or 600 ms). For both durations, minimum audible "movement" angles ("MAMA's") were on the order of 5 degrees for stimuli presented at 0 degrees azimuth (straight ahead), and increased to greater than 30 degrees for stimuli presented at +/- 90 degrees azimuth. Experiment 2 further investigated MAMA's at 0 degrees azimuth, employing two different procedures to track threshold: holding stimulus duration constant (at 100-600 ms) while varying velocity; or holding the velocity constant (at 22 degrees-360 degrees/s) while varying duration. Results from the two procedures agreed with each other and with the MAMA's determined by Perrott and Musicant for actually moving sound sources [J. Acoust. Soc. Am. 62, 1463-1466 (1977b)]: As stimulus duration decreased below 100-150 ms, the MAMA's increased sharply from 5 degrees-20 degrees or more, indicating that there is some minimum integration time required for subjects to perform optimally in an auditory spatial resolution task. Experiment 3 determined differential "velocity" thresholds employing simulated reference velocities of 0 degrees-150 degrees/s and stimulus durations of 150-600 ms. As with experiments 1 and 2, the data are more easily summarized by considering angular distance than velocity: For a given "extent of movement" of a reference target, about 4 degrees-10 degrees additional extent is required for threshold discrimination between two "moving" targets, more or less independently of stimulus duration or reference velocity. These data suggest that for the range of simulated velocities employed in these experiments, subjects respond to spatial changes--not velocity per se--when presented with a "motion" detection or discrimination task.     

Resolution in azimuth sound localization in the Mongolian gerbil (Meriones unguiculatus)

Minimum resolvable angles (MRAs) for sound localization in azimuth in the gerbil were determined in a behavioral study using tones, 300-Hz bands of noise centered at frequencies between 500 Hz and 8 kHz and broad-band noise of on average 60 dB SPL overall level. Using the method of constant stimuli, seven gerbils were trained in a two-alternative-forced-choice procedure to indicate if sounds were presented to them from the left or from the right by choosing the left or right arm of a Y-shaped cage. The MRA is the minimum angle between two loudspeaker locations that the gerbils discriminated. Animals were either stimulated from the front (N=4) or from the back (N=3). The MRA for broad-band noise randomly varying in level by +/- 6 dB was 23 degrees and 45 degrees for gerbils stimulated from the front or back, respectively. Generally a gerbil's MRA for tones declined up to 2 kHz reaching 20 degrees and 31 degrees for gerbils stimulated from the front or back, respectively, and the MRA was generally increased above this frequency. Results for narrow-band noise stimuli were similar. Results are discussed with respect to the available interaural cues and physiological mechanisms of sound localization in the gerbil.     

Minimum audible angle thresholds obtained under conditions in which the precedence effect is assumed to operate

Two experiments were conducted to examine the ability of human listeners to localize the "lag" or "echo" source in a precedence effect paradigm. A 5-ms noise burst was presented from a source located between 554-279 cm from the subject. This "lead" source was always located at 0 degrees azimuth. At the same time, one of two sources located at a distance of 610 cm from the subject was also activated with the same 5-ms noise burst. The subject's task was to identify which lag source had been active. Across sessions, the angular distance between the lag sources was varied, so as to allow a determination of the minimum audible angle (MAA) that could be resolved. Tests were run in a room designed to minimize reflections and in a hallway that was acoustically quite complex. No systematic differences in MAA thresholds were observed as a function of the environment employed. MAA thresholds obtained without the signal from the lead speaker were less than 1 degree for four of the five subjects tested. The precedence effect, as measured by the change in the MAA threshold, appears to have only a modest influence on localization performance. Under conditions in which the lead source was concurrently active, the thresholds were generally elevated by only 2 degrees-4 degrees. A reduction of this magnitude in the ability to resolve the position of the lag source does not seem to be sufficient, in itself, to account for the excellent localization performance frequently observed in reflective environments.     

Frequency resolution for diotic and dichotic listening conditions compared using the bandlimiting measure and a modified bandlimiting measure

Two experiments were performed that examined the relation between frequency selectivity for diotic and dichotic stimuli. Subjects were eight normal-hearing listeners. In each experiment, a 500-Hz pure tone of 400-ms duration was presented in continuous noise. In the diotic listening conditions, a signal and noise were presented binaurally with no interaural differences (So and No, respectively). In the dichotic listening conditions, the signal or noise at one ear was 180 degrees out-of-phase relative to the respective stimulus at the other ear (S pi and N pi, respectively). The first experiment examined frequency selectivity using the bandlimiting measure. Here, signal thresholds were determined as a function of masker bandwidth (50, 100, 250, 500, and 1000 Hz) for SoNo, S pi No, and SoN pi listening conditions. The second experiment used a modified bandlimiting measure. Here, signal thresholds (So and S pi) were determined with a relatively narrow No band of masker energy (50 Hz wide) centered about the signal. Then, a second No narrow-band masker (30 Hz wide) was added at another frequency region, and signal thresholds were reestablished. The results of the two experiments indicated that listeners process a wider band of frequencies when resolving dichotic stimuli than when resolving diotic or monotic stimuli. The results also indicated that the bandlimiting measure may underestimate the spectral band processed upon dichotic stimulation. Results are interpreted in terms of an across-ear and across-frequency processing of waveform amplitude envelope.     

Auditory brainstem response recovery in the dolphin as revealed by double sound pulses of different frequencies.

Recovery of auditory brainstem responses (ABR) in a bottlenose dolphin was studied in conditions of double-pip stimulation when two stimuli in a pair differed in frequency and intensity. When the conditioning and test stimuli were of equal frequencies, the test response was markedly suppressed at short interstimulus intervals; complete recovery appeared at intervals from about 2 ms (when two stimuli were of equal intensity) to 10-20 ms (when the conditioning stimulus exceeded the test by up to 40 dB). When the two stimuli were of different frequencies, the suppression diminished and was almost absent at a half-octave difference even if the conditioning stimulus exceeded the test one by 40 dB. Frequency-dependence curves (ABR amplitude dependence on frequency difference between the two stimuli) had equivalent rectangular bandwidth from +/-0.2 oct at test stimuli of 20 dB above threshold to +/-0.5 oct at test stimuli of 50 dB above threshold.     

Discrimination of sound source velocity in human listeners

The ability of six human subjects to discriminate the velocity of moving sound sources was examined using broadband stimuli presented in virtual auditory space. Subjects were presented with two successive stimuli moving in the frontal horizontal plane level with the ears, and were required to judge which moved the fastest. Discrimination thresholds were calculated for reference velocities of 15, 30, and 60 degrees/s under three stimulus conditions. In one condition, stimuli were centered on 0 degrees azimuth and their duration varied randomly to prevent subjects from using displacement as an indicator of velocity. Performance varied between subjects giving median thresholds of 5.5, 9.1, and 14.8 degrees/s for the three reference velocities, respectively. In a second condition, pairs of stimuli were presented for a constant duration and subjects would have been able to use displacement to assist their judgment as faster stimuli traveled further. It was found that thresholds decreased significantly for all velocities (3.8, 7.1, and 9.8 degrees/s), suggesting that the subjects were using the additional displacement cue. The third condition differed from the second in that the stimuli were "anchored" on the same starting location rather than centered on the midline, thus doubling the spatial offset between stimulus endpoints. Subjects showed the lowest thresholds in this condition (2.9, 4.0, and 7.0 degrees/s). The results suggested that the auditory system is sensitive to velocity per se, but velocity comparisons are greatly aided if displacement cues are present.     

Auditory intensity perception: successive versus simultaneous, across-channel discriminations.

This study measures the ability of observers to compare the intensities of two stimuli occupying different frequency regions. It includes three experiments, each experiment having two conditions. In one condition, the two stimuli to be compared were presented simultaneously within each interval; this condition has been called profile analysis. In the other condition, the two stimuli were presented successively within each interval. Because the overall level of the stimuli was randomized between intervals, the observers were encouraged to compare the intensities of the two stimuli within each observation interval rather than between intervals. The stimuli were two simple tones in experiment 1 and two tonal complexes in both experiments 2 and 3. The stimuli used in experiments 2 and 3 differed in frequency. The results show that simultaneous comparisons are superior to successive comparisons. For simple tones, the difference in threshold is about 8 dB; for complexes with 10 to 11 components, the difference in threshold is about 15 dB. These differences can be explained by assuming that internal noises in different channels were partially correlated when stimuli in those channels were presented simultaneously and were independent when the stimuli were presented successively. Cancellation of the correlated noise is therefore possible with simultaneous comparisons, making such discrimination better than that achievable with successive comparisons.     

Mechanisms determining the salience of coloration in echoed sound: influence of interaural time and level differences

This study investigates whether the salience of the pitch associated with a single reflection of a broadband sound, such as noise, is determined by the monaural information mediated by the stimuli at the two ears, or by the relative locations of the primary sound and the reflection. Pitch strength was measured as a function of the reflection delay and the lateral displacement between the primary sound and the reflection. Thereby, lateral displacement was produced by means of interaural time differences (ITDs) in experiment 1 and interaural level differences (ILDs) in experiment 3. The results from both experiments are in accordance with the assumption that the strength of the pitch associated with a reflection is based on a central average of the internal representations of the stimuli at the two ears. This notion was corroborated by experiment 2, which showed that the results from experiment 1 could be mimicked by simply adding the stimuli from the two ears and presenting the merged stimulus identically to both ears.     

Localization cues with bilateral cochlear implants

Selected subjects with bilateral cochlear implants (CIs) showed excellent horizontal localization of wide-band sounds in previous studies. The current study investigated localization cues used by two bilateral CI subjects with outstanding localization ability. The first experiment studied localization for sounds of different spectral and temporal composition in the free field. Localization of wide-band noise was unaffected by envelope pulsation, suggesting that envelope-interaural time difference (ITD) cues contributed little. Low-pass noise was not localizable for one subject and localization depended on the cutoff frequency for the other which suggests that ITDs played only a limited role. High-pass noise with slow envelope changes could be localized, in line with contribution of interaural level differences (ILDs). In experiment 2, processors of one subject were raised above the head to void the head shadow. If they were spaced at ear distance, ITDs allowed discrimination of left from right for a pulsed wide-band noise. Good localization was observed with a head-sized cardboard inserted between processors, showing the reliance on ILDs. Experiment 3 investigated localization in virtual space with manipulated ILDs and ITDs. Localization shifted predominantly for offsets in ILDs, even for pulsed high-pass noise. This confirms that envelope ITDs contributed little and that localization with bilateral CIs was dominated by ILDs.     

Two-dimensional sound localization by human listeners

This study measured the ability of subjects to localize broadband sound sources that varied in both horizontal and vertical location. Brief (150 ms) sounds were presented in a free field, and subjects reported the apparent stimulus location by turning to face the sound source; head orientation was measured electromagnetically. Localization of continuous sounds also was tested to estimate errors in the motor act of orienting with the head. Localization performance was excellent for brief sounds presented in front of the subject. The smallest errors, averaged across subjects, were about 2 degrees and 3.5 degrees in the horizontal and vertical dimensions, respectively. The sizes of errors increased, for more peripheral stimulus locations, to maxima of about 20 degrees. Localization performance was better in the horizontal than in the vertical dimension for stimuli located on or near the frontal midline, but the opposite was true for most stimuli located further peripheral. Front/back confusions occurred in 6% of trials; the characteristics of those responses suggest that subjects derived horizontal localization information principally from interaural difference cues. The generally high level of performance obtained with the head orientation technique argues for its utility in continuing studies of sound localization.     

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.     

Similar patterns of learning and performance variability for human discrimination of interaural time differences at high and low frequencies

Sound source localization on the horizontal plane is primarily determined by interaural time differences (ITDs) for low-frequency stimuli and by interaural level differences (ILDs) for high-frequency stimuli, but ITDs in high-frequency complex stimuli can also be used for localization. Of interest here is the relationship between the processing of high-frequency ITDs and that of low-frequency ITDs and high-frequency ILDs. A few similarities in human performance with high- and low-frequency ITDs have been taken as evidence for similar ITD processing across frequency regions. However, such similarities, unless accompanied by differences between ITD and ILD performance on the same measure, could potentially reflect processing attributes common to both ITDs and ILDs rather than to ITDs only. In the present experiment, both learning and variability patterns in human discrimination of ITDs in high-frequency amplitude-modulated tones were examined and compared to previously obtained data with low-frequency ITDs and high-frequency ILDs. Both patterns for high-frequency ITDs were more similar to those for low-frequency ITDs than for high-frequency ILDs. These results thus add to the evidence supporting similar ITD processing across frequency regions, and further suggest that both high- and low-frequency ITD processing is less modifiable and more noisy than ILD processing.     

Comparison of target detection capabilities of the beluga and bottlenose dolphin

The echolocation detection capabilities of a beluga (Delphinapterus leucas) and an Atlantic bottlenose dolphin (Tursiops truncatus) were directly compared in a target detection experiment. Both animals were trained to detect targets in the presence of masking noise. Targets were stainless-steel, water-filled spheres 7.62 and 22.86 cm in diameter. Target ranges of 16.5 and 40 m were used with the 7.62-cm sphere and 80 m with the 22.86-cm sphere. Masking noise with a flat spectrum from 40-160 kHz was projected from a spherical transducer placed 4 or 5 m, depending on the target distance, from the animal hoop station in line with the target. Target detection performance was determined as a function of masking noise level at each target range. The echo-to-noise ratio (Ee/No)max for the beluga at the 75% correct response threshold was approximately 1.0 dB compared to about 10 dB for the dolphin. The differences of each animal's detection performance across the three ranges were consistent with target strength and transmission loss differences. It is speculated that the difference in performance between the two species may be due to differences in critical bandwidth, signal processing capability, or echolocation strategy.     

Simultaneously measured behavioral and electrophysiological hearing thresholds in a bottlenose dolphin (Tursiops truncatus)

Dolphin auditory thresholds obtained via evoked potential audiometry may deviate from behavioral estimates by 20 dB or more. Differences in the sound source, stimulus presentation method, wave form, and duration may partially explain these discrepancies. To determine the agreement between behavioral and auditory evoked potential (AEP) threshold estimates when these parameters are held constant, behavioral and AEP hearing tests were simultaneously conducted in a bottlenose dolphin. Measurements were made in-air, using sinusoidal amplitude-modulated tones continuously projected via a transducer coupled to the pan region of the dolphin's lower jaw. Tone trials were presented using the method of constant stimuli. Behavioral thresholds were estimated using a 50% correct detection. AEP thresholds were based on the envelope following response and 50% correct detection. Differences between AEP and behavioral thresholds were within +/-5 dB, except at 10 kHz (12 dB), 20 kHz (8 dB), 30 kHz (7 dB), and 150 kHz (24 dB). In general, behavioral thresholds were slightly lower, though this trend was not significant. The results demonstrate that when the test environment, sound source, stimulus wave form, duration, presentation method, and analysis are consistent, the magnitude of the differences between AEP and behavioral thresholds is substantially reduced.     

Fidelity of three-dimensional-sound reproduction using a virtual auditory display

The fidelity of reproducing free-field sounds using a virtual auditory display was investigated in two experiments. In the first experiment, listeners directly compared stimuli from an actual loudspeaker in the free field with those from small headphones placed in front of the ears. Headphone stimuli were filtered using head-related transfer functions (HRTFs), recorded while listeners were wearing the headphones, in order to reproduce the pressure signatures of the free-field sounds at the eardrum. Discriminability was investigated for six sound-source positions using broadband noise as a stimulus. The results show that the acoustic percepts of real and virtual sounds were identical. In the second experiment, discrimination between virtual sounds generated with measured and interpolated HRTFs was investigated. Interpolation was performed using HRTFs measured for loudspeaker positions with different spatial resolutions. Broadband noise bursts with flat and scrambled spectra were used as stimuli. The results indicate that, for a spatial resolution of about 6 degrees, the interpolation does not introduce audible cues. For resolutions of 20 degrees or more, the interpolation introduces audible cues related to timbre and position. For intermediate resolutions (10 degrees - 15 degrees) the data suggest that only timbre cues were used.     

Assessing temporary threshold shift in a bottlenose dolphin (Tursiops truncatus) using multiple simultaneous auditory evoked potentials

Hearing sensitivity was measured in a bottlenose dolphin before and after exposure to an intense 20-kHz fatiguing tone in three different experiments. In each experiment, hearing was characterized using both the auditory steady-state response (ASSR) and behavioral methods. In experiments 1 and 2, ASSR stimuli consisted of seven frequency-modulated tones, each with a unique carrier and modulation frequency. The tones were simultaneously presented to the subject and the ASSR at each modulation rate measured to determine the effects of the sound exposure at the corresponding carrier frequency. In experiment 3 behavioral thresholds and ASSR input-output functions were measured at a single frequency before and after three exposures. Hearing loss was frequency-dependent, with the largest temporary threshold shifts occurring (in order) at 30, 40, and 20 kHz. ASSR threshold shifts reached 40-45 dB and were always larger than behavioral shifts (19-33 dB). The ASSR input-output functions were represented as the sum of two processes: a low threshold, saturating process and a higher threshold, linear process, that react and recover to fatigue at different rates. The loss of the near-threshold saturating process after exposure may explain the discrepancies between the ASSR and behavioral threshold shifts.     

The effect of aging on horizontal plane sound localization

An experiment was conducted to determine the effect of aging on sound localization. Seven groups of 16 subjects, aged 10-81 years, were tested. Sound localization was assessed using six different arrays of four or eight loudspeakers that surrounded the subject in the horizontal plane, at a distance of 1 m. For two 4-speaker arrays, one loudspeaker was positioned in each spatial quadrant, on either side of the midline or the interaural axis, respectively. For four 8-speaker arrays, two loudspeakers were positioned in each quadrant, one close to the midline and the second separated from the first by 15 degrees, 30 degrees, 45 degrees, or 60 degrees. Three different 300-ms stimuli were localized: two one-third-octave noise bands, centered at 0.5 and 4 kHz, and broadband noise. The stimulus level (75 dB SPL) was well above hearing threshold for all subjects tested. Over the age range studied, percent-correct sound-source identification judgments decreased by 12%-15%. Performance decrements were apparent as early as the third decade of life. Broadband noise was easiest to localize (both binaural and spectral cues were available), and the 0.5-kHz noise band, the most difficult to localize (primarily interaural temporal difference cue available). Accuracy was relatively higher in front of than behind the head, and errors were largely front/back mirror image reversals. A left-sided superiority was evident until the fifth decade of life. The results support the conclusions that the processing of spectral information becomes progressively less efficient with aging, and is generally worse for sources on the right side of space.     

Concurrent minimum audible angle: a re-examination of the concept of auditory spatial acuity

Minimum audible angle was measured for simultaneous acoustic events. Localization of concurrent events was found to be a direct function of the spectral differences between the events, the angle between the sources, and the location of the sources within the field defined by the subject. In the latter case, the m.a.a. was smallest with sources placed symmetrically about the listener's median plane and maximal at the extreme lateral portions. Post-hoc tests were completed which indicate that the spectral limits for concurrent localization is dependent both upon the angular separation of the sources and the position within the field as defined by the locus of the subject. The functions obtained approach the values reported by Mills [J. Acoust. Soc. Am. 30, 237-246(1958)] as the temporal overlap between the concurrent events decreased. The present results suggest that a single localization function may exist with the optimal performance observed with fully successive stimuli and poorest performance in the condition involving simultaneous events. The implications of these results are discussed.     

Lateralization of acoustic signals by dichotically listening budgerigars (Melopsittacus undulatus)

Sound localization allows humans and animals to determine the direction of objects to seek or avoid and indicates the appropriate position to direct visual attention. Interaural time differences (ITDs) and interaural level differences (ILDs) are two primary cues that humans use to localize or lateralize sound sources. There is limited information about behavioral cue sensitivity in animals, especially animals with poor sound localization acuity and small heads, like budgerigars. ITD and ILD thresholds were measured behaviorally in dichotically listening budgerigars equipped with headphones in an identification task. Budgerigars were less sensitive than humans and cats, and more similar to rabbits, barn owls, and monkeys, in their abilities to lateralize dichotic signals. Threshold ITDs were relatively constant for pure tones below 4 kHz, and were immeasurable at higher frequencies. Threshold ILDs were relatively constant over a wide range of frequencies, similar to humans. Thresholds in both experiments were best for broadband noise stimuli. These lateralization results are generally consistent with the free field localization abilities of these birds, and add support to the idea that budgerigars may be able to enhance their cues to directional hearing (e.g., via connected interaural pathways) beyond what would be expected based on head size.     

Acoustic pursuit of invisible moving targets by cats.

Head movements evoked by an invisible acoustic target were used as a metric to analyze localization of moving sources of sound in naive cats. The target was presented in the lateral sound field and moved along an arc at constant angular speeds. Head-movement trajectories were characterized by a large-magnitude orienting component that undershot the target, and a tracking component elicited by the target during acoustic pursuit. The tracking component was characterized by a succession of stepwise head movements that maintained a relatively close alignment of the median plane of the head with the moving acoustic target.