Localization of sound in the vertical plane with and without high-frequency spectral cues.

Binaural localization of 3.0-kHz high- and lowpass noise presented in the median vertical plane (MVP) and lateral vertical plane (LVP) was investigated. We anticipated superior performance when localizing the highpass noise by virtue of the availability of pinna cues. The viability of this supposition was strengthened by monaural localization tests in which performance proficiency for the highpass noise exceeded that for the lowpass noise (p less than .01). The main result showed that binaural localization of proficiency for highpass noise surpassed that for lowpass noise for all listening conditions (p less than .01). However, the importance of binaural temporal and level differences in vertical-plane localization was demonstrated by the highly respectable performances when the lowpass noise was presented in the LVP. Data from binaural localization in the MVP and monaural localization in the LVP suggested that the influence of pinna cues diminishes for source elevations above 45 degrees.     

Binaural and monaural localization of sound in two-dimensional space

Two experiments were conducted. In experiment 1, part 1, binaural and monaural localization of sounds originating in the left hemifield was investigated. 104 loudspeakers were arranged in a 13 x 8 matrix with 15 degrees separating adjacent loudspeakers in each column and in each row. In the horizontal plane (HP), the loudspeakers extended from 0 degrees to 180 degrees; in the vertical plane (VP), they extended from -45 degrees to 60 degrees with respect to the interaural axis. Findings of special interest were: (i) binaural listeners identified the VP coordinate of the sound source more accurately than did monaural listeners, and (ii) monaural listeners identified the VP coordinate of the sound source more accurately than its HP coordinate. In part 2, it was found that foreknowledge of the HP coordinate of the sound source aided monaural listeners in identifying its VP coordinate, but the converse did not hold. In experiment 2, part 1, localization performances were evaluated when the sound originated from consecutive 45 degrees segments of the HP, with the VP segments extending from -22.5 degrees to 22.5 degrees. Part 2 consisted of measuring, on the same subjects, head-related transfer functions by means of a miniature microphone placed at the entrance of their external ear canal. From these data, the 'covert' peaks (defined and illustrated in text) of the sound spectrum were extracted. This spectral cue was advanced to explain why monaural listeners in this study as well as in other studies performed better when locating VP-positioned sounds than when locating HP-positioned sounds. It is not claimed that there is inherent advantage for localizing sound in the VP; rather, monaural localization proficiency, whether in the VP or HP, depends on the availability of covert peaks which, in turn, rests on the spatial arrangement of the sound sources.     

Localization of sound in the vertical plane with and without high-frequency spectral cues

Binaural localization of 3.0-kHz high- and lowpass noise presented in the median vertical plane (MVP) and lateral vertical plane (LVP) was investigated. We anticipated superior performance when localizing the highpass noise hy virtue of the availability of pinna cues. The viability of this supposition was strengthened by monaural localization tests in which performance proficiency for the highpass noise exceeded that for the lowpass noise (p < .01). The main result showed that binaural localization of proficiency for highpass noise surpassed that for lowpass noise for all listening conditions (p < .01). However, the importance of binaural temporal and level differences in vertical-plane localization was demonstrated by the highly respectable performances when the lowpass noise was presented in the LVP. Data from binaural localization in the MVP and monaural localization in the LVP suggested that the influence of pinna cues diminishes for source elevations above 45°.     

Auditory apparent motion under binaural and monaural listening conditions

This investigation examined the ability of listeners to perceive apparent motion under binaural and monaural listening conditions. Fifty-millisecond broadband noise sources were presented through two speakers separated in space by either 10 degrees, 40 degrees, or 160 degrees, centered about the subject's midline. On each trial, the sources were temporally separated by 1 of 12 interstimulus onset intervals (ISOIs). Six listeners were asked to place their experience of these sounds into one of five categories (single sound, simultaneous sounds, continuous motion, broken motion, or successive sounds), and to indicate either the proper temporal sequence of presentation or the direction of motion, depending on whether or not motion was perceived. Each listener was tested at all spatial separations under binaural and monaural listening conditions. Motion was perceived in the binaural listening condition at all spatial separations tested for ISOIs between 20 and 130 msec. In the monaural listening condition, motion was reliably heard by all subjects at 10 degrees and 40 degrees for the same range of ISOIs. At 160 degrees, only 3 of the 6 subjects consistently reported motion. However, when motion was perceived in the monaural condition, the direction of motion could not be determined.     

The effects of attenuation of frequency segments on binaural localization of sound

Perceived location of tonal stimuli d narrow noise bands presented in two-dimensional space varies in an orderly manner with changes in stimulus frequency. Hence, frequency has a referent in space that is most apparent during monaural listening. The assumption underlying the present study is that maximum sound pressure level measured at the ear canal entrance for the various frequencies serves as a prominent spectral cue for their spatial referents. Even in binaural localization, location judgments in the vertical plane are strongly influenced by spatial referents. We measured sound pressure levels at the left ear canal entrance for 1.0-kHz-wide noise bands, centered from 4.0 kHz through 10.0 kHz, presented at locations from 60° through ?45° in the vertical plane; the horizontal plane coordinate was fixed at ?90°. On the basis of these measurements, we fabricated three different band-stop stimuli in which differently centered 2.0-kHz-wide frequency segments were filtered from a broadband noise. Unfiltered broadband noise served as the remaining stimulus. Localization accuracy differed significantly among stimulus conditions (p<.01). Where in the vertical plane most errors were made depended on which frequency segment was filtered from the broadband noise.     

The spatial attributes of stimulus frequency and their role in monaural localization of sound in the horizontal plane

Listeners, whose right ears were blocked, located low-intensity sounds originating from loudspeakers placed 15 deg apart along the horizontal plane on the side of the open, or functioning, ear. In Experiment 1, the stimuli consisted of noise bursts, 1.0 kHz wide and centered at 4.0 through 14.0 kHz in steps of .5 kHz. We found that the apparent location of the noise bursts was governed by their frequency composition. Specifically, as the center frequency was increased from 4.0 to about 8.0 kHz, the sound appeared to move away from the frontal sector and toward the side. This migration pattern of the apparent sound source was observed again when the center frequency was increased from 8.0 to about 12.0 kHz. Then, with center frequencies of 13.0 and 14.0 kHz, the sound appeared once more in front. We referred to this relation between frequency composition and apparent location in terms of spatial referent maps. In Experiment 2, we showed that localization was more proficient if the frequency content of the stimulus served to connect adjacent spatial referent maps rather than falling within a single map. By these means, we have further elucidated the spectral cues utilized in monaural localization of sound in the horizontal plane.     

Spectral cues which influence monaural Localization in the horizontal plane

An extensive series of behavioral tests was carried out to determine what region, or regions, of the sound spectrum were critical for locating sounds monaurally in the horizontal plane. Seven subjects were requested to locate narrow bands of noise centered at different frequencies, combinations of these noise bands, low-pass, high-pass, and broadband noise. As observed in an earlier study, increasing bandwidth did not necessarily lead to improved localization performance until the band became broad, including, for example, all frequencies above 4.0 kHz. What seems to be happening is that listeners perceive narrow bands of noise originating from restricted places in the horizontal plane which may differ one from another depending on the frequency composition of the stimulus. In several instances, if two noise bands were presented simultaneously, the resulting stimulus was located with reasonable accuracy provided each component, when presented singly, was perceived as emanating from clearly separate azimuthal positions. If, however, two noise bands, which were perceived to originate from approximately the same azimuthal position when presented singly, were now presented simultaneously, the resulting stimulus still was perceived to originate from the same region of the horizontal plane. This, then, is a case where augmenting the spectral content of the stimulus does not bring about improved performance. We suggest that the expression of judgmental biases in the apparent location of a band of noise may prove useful for understanding why some stimuli of specified width and center frequency are localizable while others are not.     

Auditory spatial responses of young guinea pigs (Cavia porcellus) during and after ear blocking

Newborn guinea pigs were tested to determine their ability to approach an auditory stimulus early in development. Observations of the behavior of 1-4-day-old animals in a circular eight-choice maze revealed a pronounced tendency to orient toward and approach a tape-recorded signal of guinea pig vocalizations. The occurrence of approach responses was reduced to chance in animals tested with one ear occluded by wax ear plugs which attenuated but did not totally eliminate sound. The effect of monaural ear blocks was more severe than binaural blocks, which reflects the importance of binaural cues in the maintenance of approach responses to sound. In a second study, the ability of older animals, 11-31 days of age, was examined. Directional approach responses to sound were also evident at this age, and ear plugs disrupted performance only under monaural conditions. Furthermore, in animals raised from birth with monaural ear blocks but tested without ear plugs, there was a subsequent disruption of performance for at least 21 days. These results indicate the importance of binaural cues in the development of early auditory spatial responses and suggest the need for appropriate binaural experience for subsequent localization of sounds.     

Acuity of sound localisation: a topography of auditory space. III. Monaural hearing conditions

A study is reported in which the acuity of azimuth and elevation discrimination under monaural listening conditions was measured. Six subjects localised a sound source (white noise through a speaker) which varied in position over a range of elevations (-40 degrees to +40 degrees) and azimuths (0 degrees to 180 degrees), at 10 degrees intervals, on the left side of the head. Monaural listening conditions were established by the fitting of an ear defender and one earmuff to the right ear. The absolute and algebraic, azimuth and elevation errors were measured for all subjects at each position of the source. The results indicate that all subjects suffered a marked reduction of azimuth acuity under monaural conditions, although a coarse capacity to discriminate azimuth still remained. Considerable between-subject variability was observed. Front/back discrimination was retained, although it was slightly impaired compared to that observed under normal listening conditions. Elevation discrimination was, on the whole, quite good under monaural conditions. However, a comparison of the performance of these subjects under monaural conditions with that observed under normal listening conditions indicated that some reduction in elevation localisation acuity occurred in the frontal quadrants in the median plane and in the upper quadrants of more lateral source positions. The reduction in acuity seen in these regions is attributed to the loss of information from the pinna of the occluded ear rather than to the observed reduction in azimuth error. The results provide partial support for the binaural pinna disparity model.     

The linkage between stimulus frequency and covert peak areas as it relates to monaural localization

Head-related transfer functions for differently centered narrow noise bands were obtained on 6 subjects. Derived from these measurements were covert peak areas (CPAs), defined as the spatial constellation of loudspeakers that generates maximal sound pressure at the entrance of the ear canal for specific bands of frequency. On the basis of previous data, we proposed that different frequency bands served as important spectral cues for monaural localization of sounds from different loci and that location judgments were directed toward the CPAs associated with the different bands. In the first study, the stimuli were bandpass filtered so that they contained only those frequencies whose associated CPAs occupied either the monaural listener’s “upper” or “lower” spatial regions. Loudspeakers, separated by 15°, were stationed in the left hemifield, ranging from 0° to 180° azimuth and ?45° to 60° elevation. Subjects reported the loudspeaker from which the sound appeared to originate. Judgments of the sound’s elevation were in general accord with the CPAs associated with the different frequency segments. In the second study, monaural localization tests were administered in which different 2.0-kHz-wide frequency bands linked with specific CPAs were notch filtered from a 3.5-kHz highpass noise band. For the control condition, the highpass noise was unfiltered. The data demonstrated that filtering a frequency segment linked with specific CPAs resulted in significantly fewer location responses directed toward that particular spatial region. These results demonstrate in greater detail the relation between the directional filtering properties of the pinna and monaural localization of sound.     

Frequency-shift detectors bind binaural as well as monaural frequency representations

Previous psychophysical work provided evidence for the existence of automatic frequency-shift detectors (FSDs) that establish perceptual links between successive sounds. In this study, we investigated the characteristics of the FSDs with respect to the binaural system. Listeners were presented with sound sequences consisting of a chord of pure tones followed by a single test tone. Two tasks were performed. In the "present/absent" task, the test tone was either identical to one of the chord components or positioned halfway in frequency between two components, and listeners had to discriminate between these two possibilities. In the "up/down" task, the test tone was slightly different in frequency from one of the chord components and listeners had to identify the direction (up or down) of the corresponding shift. When the test tone was a pure tone presented monaurally, either to the same ear as the chord or to the opposite ear, listeners performed the up/down task better than the present/absent task. This paradoxical advantage for directional frequency shifts, providing evidence for FSDs, persisted when the test tone was replaced by a dichotic stimulus consisting of noise but evoking a pitch sensation as a consequence of binaural processing. Performance in the up/down task was similar for the dichotic stimulus and for a monaural narrow-band noise matched in pitch salience to it. Our results indicate that the FSDs are insensitive to sound localization mechanisms and operate on central frequency representations, at or above the level of convergence of the monaural auditory pathways.     

The linkage between stimulus frequency and covert peak areas as it relates to monaural localization.

Head-related transfer functions for differently centered narrow noise bands were obtained on 6 subjects. Derived from these measurements were covert peak areas (CPAs), defined as the spatial constellation of loudspeakers that generates maximal sound pressure at the entrance of the ear canal for specific bands of frequency. On the basis of previous data, we proposed that different frequency bands served as important spectral cues for monaural localization of sounds from different loci and that location judgments were directed toward the CPAs associated with the different bands. In the first study, the stimuli were bandpass filtered so that they contained only those frequencies whose associated CPAs occupied either the monaural listener's "upper" or "lower" spatial regions. Loudspeakers, separated by 15 degrees, were stationed in the left hemifield, ranging from 0 degree to 180 degrees azimuth and -45 degrees to 60 degrees elevation. Subjects reported the loudspeaker from which the sound appeared to originate. Judgments of the sound's elevation were in general accord with the CPAs associated with the different frequency segments. In the second study, monaural localization tests were administered in which different 2.0-kHz-wide frequency bands linked with specific CPAs were notch filtered from a 3.5-kHz highpass noise band. For the control condition, the highpass noise was unfiltered. The data demonstrated that filtering a frequency segment linked with specific CPAs resulted in significantly fewer location responses directed toward that particular spatial region. These results demonstrate in greater detail the relation between the directional filtering properties of the pinna and monaural localization of sound.     

Control of responding by the location of sound: role of binaural cues.

In auditory localization experiments, where the subject observes from a fixed position, both relative sound intensity and arrival time at the two ears determine the extent of localization performance. The present experiment investigated the role of binaural cues in a different context, the sound-position discrimination task, where the subject is free to move and interact with the sound source. The role of binaural cues was investigated in rats by producing an interaural imbalance through unilateral removal of the middle auditory ossicle (incus) prior to discrimination training. Discrete trial go-right/go-left sound-position discrimination of unilaterally incudectomised rats was then compared with that of normal rats and of rats with the incus of both sides removed. While bilateral incus removal affected binaural intensity and arrival times, the symmetry of sound input between the two ears was preserved. Percentage of correct responses and videotaped observations of sound approach and exploration showed that the unilateral rats failed to localize the sounding speaker. Rats with symmetrical binaural input (normal and bilaterally incudectomised rats) accurately discriminated sound position for the duration of the experiment. Previously reported monaural localization based upon following the intensity gradient to the sound source was not observed in the unilaterally incudectomised rats of the present experiment. It is concluded that sound-position discrimination depends upon the use of binaural cues.     

Development of the auditory orientation response in the albino rat (Rattus norvegicus)

The development of head orientation to auditory stimulation was examined in rat pups at Postnatal Days 8, 11, 14, 17, and 20. The animals were tested in a quiet environment with single bursts of 65 dB (SPL) broad-band noise. A reflexive head turn toward the sound was first seen on Postnatal Day 14 and subsequently on Days 17 and 20. This result demonstrates that the onset of directional auditory responses occurred between Day 11 and Day 14. The role of binaural cues in early sound orientation was examined in 17-day-old pups with monaural ligation of the external meatus. These animals were unable to localize a sound source and consistently turned toward the side of the unligated ear regardless of the position of the stimulus. Thus binaural cues were shown to be important for head orientation to sound in early development. In a separate study, head orientation to high and low frequency tone pips was examined. Directional responses were first seen on Day 12 for a 16-kHz tone and Day 14 for a 2-kHz tone. These results indicate an earlier onset for orientation to high frequency sounds in the rat.     

Monaural localization following exposure to different segments of acoustic space

Four groups of eight monaural listeners received practice on locating sounds coming from different segments of the horizontal plane prior to a test in which all sounds originated within the same region. An additional eight monaural listeners were given the final localization test without the pretest practice. Knowledge of results was withheld. The main finding was that positive transfer of training was not equally apparent for all groups. That group for which the pretest and test involved the same ear and the same azimuthal positions of loudspeakers performed best. Practice in locating rearwardly positioned sounds did not benefit the localization of frontally positioned sounds even when the same ear was functioning in both situations. Experience in locating sounds from all segments of the horizontal plane appears to be required in order to build up an adequate internal representation of the acoustic surrounds.     

Hearing in large mammals: sound-localization acuity in cattle (Bos taurus) and goats (Capra hircus).

Sound localization acuity of 3 cattle (Bos taurus) and 2 goats (Capra hircus) was determined for brief complex sounds in a two-choice procedure. Thresholds around the median sagittal plane averaged 30 degrees and 18 degrees, respectively. For comparison, thresholds were obtained in the same test apparatus for humans (0.8 degrees) and a dog (8 degrees). Although the relatively poor acuity of cattle and goats compared with most mammals comes as some surprise, given their large interaural distances and the large binaural locus cues available to them, it is not unexpected when other factors are considered. Like other poor localizers (both domesticated and nondomesticated), cattle and goats are prey species with their best vision directed throughout nearly the entire horizon. In contrast to mammals with very narrow foveal fields, they may not need very accurate locus information from their auditory systems to direct their gaze to a sound source.     

A comparison of the contrast effects in sound localization in the horizontal and vertical planes

The effect of a background sound on the auditory localization of a single sound source was examined. Nine loudspeakers were arranged crosswise in the horizontal and the median vertical plane. They ranged from -20 degrees to +20 degrees, with the center loudspeaker at 0 degree azimuth and elevation. Using vertical and horizontal centimeter scales, listeners verbally estimated the position of a 500-ms broadband noise stimulus being presented at the same time as a 2 s background sound, emitted by one of the four outer loudspeakers. When the background sound consisted of continuous broadband noise, listeners consistently shifted the apparent target positions away from the background sound locations. This auditory contrast effect, which is consistent with earlier findings, equally occurred in both planes. But when the background sound was changed to a pulse train of noise bursts, the contrast effect decreased in the horizontal plane and increased in the vertical plane. This discrepancy might be due to general differences in the processing of interaural and spectral localization information.     

Sound localization in an Old-World fruit bat (Rousettus aegyptiacus): acuity, use of binaural cues, and relationship to vision.

The passive sound-localization acuity of Egyptian fruit bats (Rousettus aegyptiacus) was determined using a conditioned-avoidance procedure. The mean minimum audible angle for left-right discrimination for 3 bats was 11.6 degrees--very near the mean for terrestrial mammals. The bats also were able to localize low- and high-frequency pure tones, indicating that they can use both binaural phase-difference and binaural intensity-difference cues to localize sound. Moreover, they were able to use the binaural phase-difference cue up to at least 5.6 kHz, which is higher than other mammals yet tested. The width of the Egyptian fruit bats' field of best vision was 27 degrees. This value is consistent with the hypothesis that the role of passive sound localization is to direct the eyes for visual scrutiny of sound sources. Thus, the passive localization abilities of these echolocating megachiropteran fruit bats do not deviate from the patterns established for nonecholocating mammals.     

Spectral information for detection of acoustic time to arrival

The exponential increase of intensity for an approaching sound source provides salient information for a listener to make judgments of time to arrival (TTA). Specifically, a listener will experience a greater rate of increasing intensity for higher than for lower frequencies during a sound source’s approach. To examine the relative importance of this spectral information, listeners were asked to make judgments about the arrival times of nine 1-octave-band sound sources (the bands were consecutive, nonoverlapping single octaves, ranging from 40–80 Hz to ~10–20 kHz). As is typical in TTA tasks, listeners tended to underestimate the arrival time of the approaching sound source. In naturally occurring and independently manipulated amplification curves, bands with center frequencies between 120 and 250 Hz caused the least underestimation, and bands with center frequencies between 2000 and 7500 Hz caused the most underestimation. This spectral influence appears to be related to the greater perceived urgency of higher-frequency sounds.     

The relative influence of pitch and timbre on the apparent location of sound in the median sagittal plane

Listeners were requested to locate sounds originating in the median sagittal plane (MSP). The stimuli, 75-Msec pulses, were repeated at a rate of 200 times/sec and were filtered to transmit narrow bands centered about 0.63, 1.6, 2.5, or 6.3 kHz. Despite the sameness of pitch generated by all stimuli, Os perceived the 0.63-, 1.6-, and 2.5-kHz-centered sounds as originating low, middle, and high, respectively, in the MSP, regardless of their actual positions. The stimuli centered about 6.3 kHz, on the other hand, were located accurately by most Os. These findings were interpreted to mean that under conditions of inadequate auditory cues, timbre, not pitch, influences perceived elevation. The implication is that timbre also served as the cue for the apparent elevation of those sounds which, due to their high-frequency components, can be located accurately.