Evoked potentials in sound localization: timing of activity along the auditory pathway.
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Acoustic Stimulation↗
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Results for “Sound Localization”
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The effect of various hearing protectors on sound localization in the horizontal and vertical planes.
Listeners were tested under free head/torso movement conditions, using a brief pulsed broadband noise signal presented from any 1 of 20 loudspeakers in two intersecting hemicircumferential arrays: one in the horizontal plane, one in the vertical. The listeners' task was to judge the whereabouts of each signal while listening in normal, open-ears conditions or while wearing various types of standard or electronic earmuffs or compressible foam earplugs. Results showed that listeners could accomplish the task successfully in normal (open-ear) conditions. Results using earplugs, nonelectronic earmuffs, and two types of dichotic (dual microphone) electronic earmuffs showed decrements in performance similar to each other. These decrements took the general form of loss of appreciation of the vertical plane and of front-rear differences in the horizontal plane. Lateral discrimination was blurred but otherwise intact. Of two diotic (single microphone) electronic earmuffs, one completely disrupted localization function and the other came close to it.
Auditory Perception↗
The effect of head rotations on vertical plane sound localization.
Current understanding gives predominant weight to stationary cues for auditory localization. Two experiments were conducted to investigate the possible existence of a dynamic cue. The first experiment involved localization of concealed sources in the upper median vertical plane (MVP) and showed, as expected, that elevation was not detectable with motionless listening when high-frequency energy was absent or when normal pinna function was distorted. Elevation under such conditions did become detectable with horizontal head rotations, provided low-frequency energy was present in the signal. This indicates that the basis of the dynamic cue is variation in the rate of transformation of low-frequency interaural time/phase differences. The second experiment involved localization of sources arrayed throughout upper and lower regions of the MVP and in the left lateral vertical plane (LVP); it showed that upper hemisphere sources can be distinguished somewhat from those in the lower hemisphere, even in motionless listening conditions, but more so with rotation. The greatest benefit for localization from rotation of the head appears to be gained for sources positioned in the front MVP.
Adult↗
The role of macaque auditory cortex in sound localization.
Bilateral ablation of auditory cortex in macaques results in both sensory and perceptual deficits. The sensory deficit is indicated by increased thresholds for left-right locus discriminations and an inability to discriminate locus within either the left of right hemifield. The perceptual deficit is indicated by the observation that the monkeys no longer appear to associate a sound with a location in space. Unilateral auditory cortex ablation results in an inability to discriminate locus within the hemifield contralateral to the lesion. It is not known whether unilateral lesions also result in a perceptual deficit.
Animals↗
The influence of pinnae-based spectral cues on sound localization.
The role of pinnae-based spectral cues was investigated by requiring listeners to locate sound, binaurally, in the horizontal plane with and without partial occlusion of their external ears. The main finding was that the high frequencies were necessary for optimal performance. When the stimulus contained the higher audio frequencies, e.g., broadband and 4.0-kHz high-pass noise, localization accuracy was significantly superior to that recorded for stimuli consisting only of the lower frequencies (4.0- and 1.0-kHz low-pass noise). This finding was attributed to the influence of the spectral cues furnished by the pinnae, for when the stimulus composition included high frequencies, pinnae occlusion resulted in a marked decline in localization accuracy. Numerous front-rear reversals occurred. Moreover, the ability to distinguish among sounds originating within the same quadrant also suffered. Performance proficiency for the low-pass stimuli was not further degraded under conditions of pinnae occlusion. In locating the 4.0-kHz high-pass noise when both, neither, or only one ear was occluded, the data demonstrated unequivocally that the pinna-based cues of the "near" ear contributed powerfully toward localization accuracy.
Auditory Perception↗
Sound localization precision under conditions of the precedence effect: effects of azimuth and standard stimuli.
Minimum audible angles (MAAs) were estimated for single noise bursts, and for burst pairs that satisfied the conditions of the precedence effect (that is, produced fused images). In one burst-pair condition, the bursts to be discriminated differed in lead location; in the other, they differed in lag location. Sounds were presented over loudspeakers. MAAs were lowest for single bursts, slightly higher for lead discrimination, and much higher for lag discrimination. Presence of a standard reference burst had no reliable effect on performance. The data are interpreted using a model of Shinn-Cunningham et al. [J. Acoust. Soc. Am. 93, 2923-2932 (1993)] in which discrimination of precedence-effect burst pairs is based on the lateral position of the auditory image, which is a weighted average of the positions of the leading and lagging bursts.
Adult↗
Sound localization in noise in hearing-impaired listeners.
The present study assesses the ability of four listeners with high-frequency, bilateral symmetrical sensorineural hearing loss to localize and detect a broadband click train in the frontal-horizontal plane, in quiet and in the presence of a white noise. The speaker array and stimuli are identical to those described by Lorenzi et al. (in press). The results show that: (1) localization performance is only slightly poorer in hearing-impaired listeners than in normal-hearing listeners when noise is at 0 deg azimuth, (2) localization performance begins to decrease at higher signal-to-noise ratios for hearing-impaired listeners than for normal-hearing listeners when noise is at +/- 90 deg azimuth, and (3) the performance of hearing-impaired listeners is less consistent when noise is at +/- 90 deg azimuth than at 0 deg azimuth. The effects of a high-frequency hearing loss were also studied by measuring the ability of normal-hearing listeners to localize the low-pass filtered version of the clicks. The data reproduce the effects of noise on three out of the four hearing-impaired listeners when noise is at 0 deg azimuth. They reproduce the effects of noise on only two out of the four hearing-impaired listeners when noise is at +/- 90 deg azimuth. The additional effects of a low-frequency hearing loss were investigated by attenuating the low-pass filtered clicks and the noise by 20 dB. The results show that attenuation does not strongly affect localization accuracy for normal-hearing listeners. Measurements of the clicks' detectability indicate that the hearing-impaired listeners who show the poorest localization accuracy also show the poorest ability to detect the clicks. The inaudibility of high frequencies, "distortions," and reduced detectability of the signal are assumed to have caused the poorer-than-normal localization accuracy for hearing-impaired listeners.
Aged↗
Progress in neurophysiology of sound localization.
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Acoustic Stimulation↗
Adaptive adjustment of unit tuning to sound localization cues in response to monaural occlusion in developing owl optic tectum.
Bimodal units in the barn owl's optic tectum are tuned to the location of auditory and visual stimuli, and are systematically organized according to their spatial tuning to form mutually aligned maps of auditory and visual space. Map alignment results from the fact that, normally, units are tuned to the values of interaural level difference (ILD) and interaural time difference (ITD) produced by a sound source at the location of their visual receptive fields (VRFs). Monaural occlusion alters the correspondence of ILD and ITD values with locations in space. We investigated the effect that raising owls with a chronic monaural occlusion has on the tuning of tectal units to ILD and ITD. Owls were monaurally occluded beginning at 1 month of age. The effects of monaural occlusion were assessed 2-4 months later by comparing the ILD and ITD tuning of units in monaurally occluded owls with the ILD and ITD tuning of units with equivalent VRFs in normal owls. ILD and ITD tuning was shifted substantially and in the direction of the unoccluded ear (the adaptive direction) in owls raised with a monaural occlusion. In most tecta, the mapped representations of ILD and ITD were shifted systematically. In addition, in some tecta, monaural occlusion induced a change in the topography of the ILD map such that ILD tuning remained essentially constant at values near 0 dB over abnormally large portions of the tectum. Across all recording sites, the average shift in ILD tuning was 9 dB (n = 396) and the average shift in ITD tuning was 40 microseconds (n = 414). In four of five animals, the magnitude of the effect was not equivalent on the two sides of the brain, the adjustments being significantly larger and more systematic on the side ipsilateral to the occlusion. Such differences in the altered ILD and ITD maps on the two sides of the brain in individual animals indicate that, although a component of the adaptive adjustment might be due to regulation of the gain and phase response of the monaural signals early in the auditory pathway, a major component of the adjustment must occur at or beyond the level where the encoding of ILDs and ITDs for left and right space separates.
Adaptation, Physiological↗
The nature and distribution of errors in sound localization by human listeners.
Measurement of localization performance will reflect errors that relate to the sensory processing of the cues to sound location and the errors associated with the method by which the subject indicates the perceived location. This study has measured the ability of human subjects to localize a short noise burst presented in the free field with the subject indicating the perceived location by pointing their nose towards the source. Subjects were first trained using a closed loop training paradigm which involved instantaneous feedback as to the accuracy of head pointing which resulted in the reduction of residual localization errors and a rapid acquisition of the task by the subjects. Once trained, 19 subjects localized between 4 and 6 blocks of 76 target locations. The data were pooled and the distribution of errors associated with each target location was examined using spherical methods. Errors in the localization estimates for about one third of the locations were rotationally symmetrical about their mean but the remaining locations were best described by an elliptical distribution (Kent distributed). For about one half of the latter locations the orientations of the directions of the greatest variance of the distributions were not aligned with the azimuth and elevation coordinates used for describing the spatial location of the targets. The accuracy (systematic errors) and the distribution of the errors (variance) in localization for our population of subjects were also examined for each test location. The size of the data set and the methods of analysis provide very reliable measures of important baseline parameters of human auditory localization.
Acoustic Stimulation↗
Proximity image effect in sound localization.
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Auditory Perception↗
Spectral cues for sound localization in cats: effects of frequency domain on minimum audible angles in the median and horizontal planes.
Rice et al. [Hear. Res. 58, 132-152 (1992)] classified directional properties of the cat's head-related transfer function (HRTF) into three frequency domains. Low frequencies (< 5 kHz) display a broad azimuth-sensitive spectral peak that establishes interaural level differences, mid frequencies (5-18 kHz) are marked with a single deep spectral notch that changes in frequency as a function of both azimuth and elevation, and high frequencies (18-50 kHz) exhibit a complex pattern of peaks and notches that shows extensive but less systematic changes with sound location. Spectral cues conveyed by the mid frequencies of broadband sounds are important in tasks that require cats to identify the actual location of acoustic stimuli [Huang and May, J. Acoust. Soc. Am. (in press)]. The present study investigates how directional cues conveyed by the mid- and high-frequency spectrum of the HRTF influence the cat's ability to discriminate between sound locations. Thresholds for spatial acuity were measured as minimum audible angles (MAAs) [Mills, J. Acoust. Soc. Am. 30, 237-246 (1958)] at positive azimuths in the interaural horizontal plane and at positive and negative elevations in the median vertical plane. The frequency domain of the noise burst had little effect on MAAs in the horizontal plane, but removal of high-frequency spectral information significantly increased thresholds at positive and negative elevations in the median plane. These results suggest that cats are sensitive to directional properties of the HRTF at frequencies above 18 kHz and may use this information to detect small changes in sound source elevation.
Animals↗
Electrophysiological evidence of a sound localizing binaural subsystem in the human auditory brainstem.
The binaural interaction components of auditory brainstem evoked potentials reflect electrical changes which are specific to binaural stimuli. Analysis of these components indicated that different click lateralizations result in spatially distinct distributions of activity in the pons, but not at more peripheral levels. The effects of ipsilateral and binaural masking on evoked activity indicated distinct binaurally- and monaurally-activated neural subsets in the human brainstem. These results on the effects of noise and the distinct distributions of pontine activity to different lateralizations provide the first electrophysiological evidence that the auditory system in the humans pons includes a subset which is specific to binaurally presented sounds and which is anatomically distributed according to the lateralization of the sound. These results suggest auditory spatial mapping, similar to other sensory systems. In contrast to other systems, in which mapping is according to receptor distribution in the periphery, auditory spatial mapping is achieved computationally at central levels of the pathway.
Acoustic Stimulation↗
Single-unit activity in the auditory cortex of monkeys actively localizing sound sources: spatial tuning and behavioral dependency.
Single-unit recordings were made from the auditory cortex of unanesthetized rhesus monkeys which were trained to perform a second localization task. Recordings were made from 196 units in 4 animals. Five sound sources were located at 0 degrees (midline) and on either side at azimuths of 37.5 degrees and 75 degrees. Almost most units responded to each of the 5 sources, 49% of the units had peak firing rates for the source on the contralateral side at 75% azimuth. Another 26% of the units had peak firing rates at the 37.5 degrees contralateral location, while only 11% had peak rates for the ipsilateral locations. In order to determine whether the behavior of actively locating a sound source affected units in auditory cortex, response rates compared under two behavioral conditions: one which required the detection of a sound regardless of location and another condition which required identification of a sound's location. of the 196 units, 16 had different response rates for the two conditions, with 15 of the 16 units having higher evoked rates in the localization task. For the 16 units, the difference in firing rate was typically observed for a single speaker location. No correlation could be determined between a unit's location within the subdivisions of auditory cortex and its spatial tuning response properties or its dependency upon behavioral condition.
Animals↗
Binaural disparity cues available to the barn owl for sound localization.
1. Bilateral recording of cochlear potentials was used to measure the variations in interaural time differences (ITDs) and interaural intensity differences (IIDs) as a free-field auditory stimulus was moved to different positions around a barn owl's head. 2. ITD varied smoothly with stimulus azimuth across a broad frequency range. 3. ITD varied minimally with stimulus elevation, except at extreme angles from the horizontal. 4. IID varied with both stimulus elevation and stimulus azimuth. Lower frequencies were more sensitive to variations in azimuth, whereas higher frequencies were more sensitive to variations in elevation. 5. The loci of spatial coordinates that form iso-IID contours and iso-ITD contours form a non-orthogonal grid that relates binaural disparity cues to sound location.
Acoustic Stimulation↗
[Sound localization in patients with asymmetrical hearing loss].
Good directional hearing ability demands good and symmetrical hearing in both ears. We report the effect of impaired hearing on the directional hearing ability of 98 patients, especially of patients with bilateral asymmetrical hearing loss. The directional testing device included 12 loudspeakers placed at 30 degree intervals in a circle with a diameter of 3.25 m, whose centre lay between the ears of the patient. In included an audiometer for producing the signals, an amplifier and a PDP11/23 computer interfaced to a loudspeaker switch bank. The subject's answers to 60 directionally randomized stimuli were recorded. During the presentation of the signal the patients were not allowed to turn their head. The patients had to name the number of the loudspeaker on the circle that they thought was producing the sound. In addition to the directional hearing test a pure-tone audiogram was done, and the middle- and high-frequency hearing loss estimated. The records of the directional hearing test were analysed in two new ways: firstly, vector analysis of the errors; secondly, averaging of the difference between the true interaural time delay and the virtual time difference, which was implicated in the possibly incorrect answer of the patient (effective delta-t-parameter). This average gives a score for the uncertainty in defining the correct "cone of confusion". In addition to the statistical analysis, two cases are reported showing the directional hearing ability of two patients with neuromas treated by transtemporal surgery, with some residual hearing.(ABSTRACT TRUNCATED AT 250 WORDS)
Audiometry↗
Single cortical neurons serve both echolocation and passive sound localization.
The pallid bat uses passive listening at low frequencies to detect and locate terrestrial prey and reserves its high-frequency echolocation for general orientation. While hunting, this bat must attend to both streams of information. These streams are processed through two parallel, functionally specialized pathways that are segregated at the level of the inferior colliculus. This report describes functionally bimodal neurons in auditory cortex that receive converging input from these two pathways. Each brain stem pathway imposes its own suite of response properties on these cortical neurons. Consequently, the neurons are bimodally tuned to low and high frequencies, and respond selectively to both noise transients used in prey detection, and downward frequency modulation (FM) sweeps used in echolocation. A novel finding is that the monaural and binaural response properties of these neurons can change as a function of the sound presented. The majority of neurons appeared binaurally inhibited when presented with noise but monaural or binaurally facilitated when presented with the echolocation pulse. Consequently, their spatial sensitivity will change, depending on whether the bat is engaged in echolocation or passive listening. These results demonstrate that the response properties of single cortical neurons can change with behavioral context and suggest that they are capable of supporting more than one behavior.
Animals↗
Sound localization, use of binaural cues and the superior olivary complex in pigs.
Noise localization thresholds and the ability to localize pure tones at 60 degrees separation were determined for three domestic pigs using a conditioned avoidance procedure. The average threshold for localizing a brief noise burst was 4.5 degrees which is much more accurate than the thresholds of other hoofed mammals, such as horses, cattle and goats. The ability of pigs to localize low-frequency tones indicates that they can use the binaural phase-difference cue. However, they were unable to localize tones of 4 kHz and higher, indicating that, like other hoofed mammals, their ability to use binaural intensity cues is greatly restricted if not completely absent. An examination of the superior olivary complex of pigs revealed that in relative size, shape and cell density it is more like that of cats than that of other hoofed mammals.
Animals↗