Gain control in the auditory system: absolute identification of intensity within and across the two ears

A number of studies have suggested that the auditory system employs nonlinear gain control to modulate its response to a broad range of stimulus intensities (e.g., Parker, Murphy, & Schneider, 2002), but the precise location and nature of this mechanism remains uncertain. To address these issues, we investigated the extent to which gain control in one ear was influenced by auditory events in the other ear. Listeners were asked to identify which of four sounds, varying in intensity (25, 30, 35, and 40 dB [SPL]), had been presented to the left ear. Subsequent test sets included an 80-dB (SPL) stimulus presented to either the left (ipsilateral) or the right (contralateral) ear. Ipsilateral presentations of the 80-dB (SPL) stimulus produced a greater reduction in identification accuracy among the original four sounds than did contralateral presentations, but both presentations caused a reduction in gain. In a second experiment, the perceived laterality of a sound was manipulated by presenting the same sound to both ears with an interaural time delay. When the 80-dB (SPL) stimulus was added to the set of quiet sounds (which were presented to both ears but were perceived to be in the left ear because of the interaural time delay), identification accuracy was reduced by the same amount, independently of whether the 80-dB (SPL) stimulus was perceived only in the ipsilateral ear or only in the contralateral ear.     

The influence of stimulus bandwidth on localization of sound in space

The ability of listeners, deprived of prominent interaural time and intensity cues, to locate noise bands differing in width was investigated. To minimize binaural cues, we placed the sound source at various positions in the median sagittal plane. To eliminate binaural cues, we occluded one ear. The stimuli consisted of broadband noise and bands of noise centered at 8.0 kHz. The width of the latter ranged from 1.0 to 6.0 kHz. The results from seven listeners showed that localization proficiency for sounds in the median sagittal plane decreased with decreases in bandwidth for both binaural and monaural listening conditions. This function was less orderly for monaural localization of horizontally positioned sounds. Another consequence of a reduction in bandwidth was an increasing tendency of listeners to select certain loudspeakers over others as the source of the sound. A previous finding showing that localization of sound in the median sagittal plane is more accurate when listening binaurally rather than monaurally was confirmed.     

An Adaptive Bias in the Perception of Looming Auditory Motion

Rising acoustic intensity can indicate movement of a sound source toward a listener. Perceptual overestimation of intensity change could provide a selective advantage by indicating that the source is closer than it actually is, providing a better opportunity for the listener to prepare for the source's arrival. In Experiment 1, listeners heard equivalent rising and falling level sounds and indicated whether one demonstrated a greater change in loudness than the other. In 2 subsequent experiments listeners heard equivalent approaching and receding sounds and indicated perceived starting and stopping points of the auditory motion. Results indicate that rising intensity changed in loudness more than equivalent falling intensity, and approaching sounds were perceived as starting and stopping closer than equidistant receding sounds. Both effects were greater for tones than for noise. Evidence is presented that suggests that an asymmetry in the neural coding of egocentric auditory motion is an adaptation that provides advanced warning of looming acoustic sources.     

Tactile "capture" of audition

Previous research has demonstrated that the localization of auditory or tactile stimuli can be biased by the simultaneous presentation of a visual stimulus from a different spatial position. We investigated whether auditory localization judgments could also be affected by the presentation of spatially displaced tactile stimuli, using a procedure designed to reveal perceptual interactions across modalities. Participants made left-right discrimination responses regarding the perceived location of sounds, which were presented either in isolation or together with tactile stimulation to the fingertips. The results demonstrate that the apparent location of a sound can be biased toward tactile stimulation when it is synchronous, but not when it is asynchronous, with the auditory event. Directing attention to the tactile modality did not increase the bias of sound localization toward synchronous tactile stimulation. These results provide the first demonstration of the tactile capture of audition.     

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.     

Tactile “capture” of audition

Previous research has demonstrated that the localization of auditory or tactile stimuli can be biased by the simultaneous presentation of a visual stimulus from a different spatial position. We investigated whether auditory localization judgments could also be affected by the presentation of spatially displaced tactile stimuli, using a procedure designed to reveal perceptual interactions across modalities. Participants made left—right discrimination responses regarding the perceived location of sounds, which were presented either in isolation or together with tactile stimulation to the fingertips. The results demonstrate that the apparent location of a sound can be biased toward tactile stimulation when it is synchronous, but not when it is asynchronous, with the auditory event. Directing attention to the tactile modality did not increase the bias of sound localization toward synchronous tactile stimulation. These results provide the first demonstration of the tactilecapture of audition.     

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.     

Auditory localization: effects of reflecting surfaces

Based on anatomical and evolutionary conceptions of the human ear, an experiment was conducted in which forty-eight human subjects were asked to localize sounds (a human voice) emitted by one of twenty-seven stationary loudspeakers in an anechoic chamber. The position of the active loudspeaker varied with respect to azimuth, distance, and elevation in three steps each. The position of a single sound-reflecting surface (about 6 m2) was varied: on the floor, on the ceiling, to the left, and to the right. The accuracy of identifying the active loudspeaker for each position of the sound-reflecting surface was compared intraindividually with the absence of reflection. The results show an overall increase in correct localizations with a sound-reflecting surface on the floor. Especially the elevation of the sound source can be detected with greater precision. Additionally, the percentage of correct localizations decreased systematically with the presence of a sound-reflecting ceiling, while the presence of sound-reflecting walls did not systematically affect the localization performance. Judgments in the horizontal plane and those of distance were not systematically influenced by the presence of a sound-reflecting surface.     

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.     

The effects of eye position and expectation on sound localization

Ss who are exposed to a sound coming from straight ahead, but who turn their eyes 20 deg to the side toward a visible speaker during the exposure period and expect to hear the sound coming from the visible source, show a shift in localization of the sound up to a maximum of about 9 deg. Ss who only turn their eyes 20 deg to the side during the exposure period show a smaller but significant shift in sound localization, while Ss who do not turn their eyes, but are led to expect that the sound will appear to come from a visible loudspeaker 20 deg to the side, show no significant shift. Comparison of test results before and after the exposure period, with eyes directed straight ahead and no visible speaker present, shows the presence of a localization aftereffect for those experimental groups that showed a significant localization shift during the exposure period. Sounds are localized a few degrees to the side of their physical location in the same direction as the shift in localization during the exposure period. Further experiments show that part, but not all, of the shift in localization during the exposure period can be understood in terms of a shift in perceived head direction. The localization aftereffects are shown not to be due to change in physical or perceived eye or head position.     

Effect of head movement on the localization of sounds in the equatorial plane

Five experiments examined the effect of head movement on the localization of sound sources in the equatorial plane. Under most conditions, head movement produces poorer localization for sound sources in the equatorial plane. Only one condition demonstrated an improvement in localization with head movement. The sound source was located toward the side of the head and the source duration was long enough to permit reorientation of the position of the head with re-spect to the sound source.     

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.     

Performance and reaction times in monaural localization of first names in the horizontal plane

Many studies have been conducted to measure monaural azimuthal sound localization performance with different sounds varying in frequency and complexity, but few have used linguistic stimuli. The present experimental design used subjects' first names in a monaural azimuthal localization task. Analysis of response accuracy showed that subjects are not more accurate in localizing their own first name than in localizing other first names and that there was no significant advantage of one ear over another. Reaction times were shorter when the subjects localized their own first name than when they localized any other first names and there was no significant ear advantage, but localizing other first names took more time with the right than with the left ear. All stimuli were better and more quickly localized on the side of the open ear, and there was no difference in acuity or velocity of localization with the two different speaker voices used. These results suggest that first names are processed through the controlateral auditory pathway and can be analyzed in the right hemisphere.     

The influence of the acoustic context on vertical sound localization in the median plane

The influence of background sounds (frames) on vertical localization of single sound sources (targets) was examined in four experiments. Loudspeakers (five targets and four frames) were positioned in the median plane, ranging from +30 degrees to -30 degrees above and below the subject's ear level. The subjects determined the vertical position of the targets by either verbal judgments or manual pointing. Frame and target sounds were presented concurrently or successively with a 1-sec interval; both consisted of (1) 300-Hz square waves, (2) noise, or (3) targets of noise and frames of 300-Hz square waves. Particularly in the second condition, the subjects consistently shifted the apparent target positions away from the frame locations. This contrast effect persisted even 1 sec after the offset of the frames. No effect was found with different waveforms for the frame and the target. Results are related to recent findings indicating a similar effect in the azimuthal dimension. Possibly the effect is based on a mechanism in which the auditory system adapts to recently heard sound source positions.     

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.     

Spatial position versus ear of entry as determinant of the auditory laterality effects: a stereophonic test.

The study concerned discriminating between ear of entry and apparent spatial position as possible determinants of lateral asymmetries in the recall of simultaneous speech messages. Apparent localization to the left or right of the median plane was created either through a time difference (.7 msec), through intensity differences between presentations of the same verbal message at the two ears, or through dichotic presentations. Right-side advantage was observed with the three types of presentation (Experiments 1, 2, and 3). The finding of right-side advantage with stereophony based on a time difference only, in the absence of intensity difference, cannot be accounted for in terms of an ear advantage and shows that apparent spatial separation of the sources can by itself produce a laterality effect. Differences in the degree of lateral asymmetry between the various conditions were also observed. The findings of Experiments 4 and 5 suggest that these differences are better explained in terms of different impressions of localization of the sound sources than in terms of relative intensity at the "privileged" ear.     

Evaluation of response methods for the localization of nearby objects

Four response methods for indicating the perceived locations of nearby objects were evaluated: the direct-location (DL) method, where a response pointer is moved directly to the perceived location of the target; the large-head (LH) and small-head (SH) methods, where the pointer is moved to the target location relative to a full-scale or half-scale manikin head; and the verbal report (VR) method, where the spherical coordinates of the target location are indicated verbally. Measurements with a visual target indicated that the DL method was relatively unbiased and considerably more accurate than the other methods, which were all roughly equivalent. Correcting for bias improved accuracy in the LH, SH, and VR responses, but not to the level of the uncorrected DL responses. Replacing the visual target with an acoustic stimulus approximately doubled the errors with the DL response but indicated similar performance in the front and rear hemispheres. The results suggest that DL is the most appropriate response method for close-range localization experiments.     

Feature mismatch effects in auditory negative priming: interference as dependent on salient aspects of prior episodes

A handful of studies have revealed that withholding a response to a sound causes impaired responding to that sound when subsequently presented (Buchner & Steffens, 2001; Mondor, Leboe, & Leboe, 2005). In the present study, we investigated whether a switch in the location of a repeated sound might represent an additional source of this auditory negative priming effect. In all three of our experiments, participants performed a series of trials in which they withheld a response to a dichotic pair of prime sounds presented to each ear. A dichotic pair of probe sounds was then presented to each ear, and the participants were required to categorize one of them. Across these experiments, the participants were slower to categorize only repeated sounds that were presented in opposite locations.     

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.     

The monaural localization of tonal stimuli

With one ear occluded, 17 listeners were asked to locate tone bursts, .25, 4, .6, .9. l.4, 2.0, 3.2, 4.8, and 7.2 kHz, generated by a loudspeaker concealed from view. The S’s response was to callout that number, from a series of numbers arranged horizontally, behind which he thought the tone bursts originated. The listeners perceived the sounds as emanating from the side of the unoccluded ear, but their judgments bore no consistent relation to the actual location of the sound source. Rather, the listeners showed a strong tendency to locate a tone burst, within the range of .9 through 7.2 kHz, in a fixed spatial relation to the next higher- and lower-pitched tone burst. Distorting the pinna of the unoccluded ear failed to modify the perceptual pattern. It was suggested that the perceived spatial relations among the various frequencies was a by-product of the tonotopic organization of the auditory nervous system.