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More accurate sound localization induced by short-term light deprivation.

Crossmodal reorganization processes in the brain are mainly associated with early blindness, on the assumption that recruitment of genuine visual areas, such as primary visual cortex, for non-visual functions results in superior auditory and tactile performance of blind, compared to sighted, humans. This study shows that in sighted subjects the accuracy of sound localization, measured by a task of head pointing to acoustic targets, is reversibly increased after short-term light deprivation of 90 min. However, only the systematic deviations from target positions (constant error) were reduced after light deprivation, while the general precision of head pointing remained unchanged. Return to pre-deprivation values was observed after 180 min of re-exposure to light. The post-deprivation change was similar, though less in magnitude, to the effect of blindness that was demonstrated previously. Generally, these findings indicate that auditory-visual crossmodal plasticity can be quite rapidly initiated by deprivation of the visual cortex from visual input. It seems possible that visual deprivation has an influence on neuronal circuits, that are involved in processing of auditory information in visual brain areas of normal sighted humans. Since exclusively the constant error in sound localization, not general performance, was changed, the present effect of visual deprivation may, however, not be attributable to reorganization processes in the sense of a compensation for the absence of vision. It is more likely that the observed change in accuracy was specifically induced by the absence of visual calibration of the neural representation of auditory space during light deprivation.

Acoustic Stimulation↗

Spectro-temporal factors in two-dimensional human sound localization.

This paper describes the effect of spectro-temporal factors on human sound localization performance in two dimensions (2D). Subjects responded with saccadic eye movements to acoustic stimuli presented in the frontal hemisphere. Both the horizontal (azimuth) and vertical (elevation) stimulus location were varied randomly. Three types of stimuli were used, having different spectro-temporal patterns, but identically shaped broadband averaged power spectra: noise bursts, frequency-modulated tones, and trains of short noise bursts. In all subjects, the elevation components of the saccadic responses varied systematically with the different temporal parameters, whereas the azimuth response components remained equally accurate for all stimulus conditions. The data show that the auditory system does not calculate a final elevation estimate from a long-term (order 100 ms) integration of sensory input. Instead, the results suggest that the auditory system may apply a "multiple-look" strategy in which the final estimate is calculated from consecutive short-term (order few ms) estimates. These findings are incorporated in a conceptual model that accounts for the data and proposes a scheme for the temporal processing of spectral sensory information into a dynamic estimate of sound elevation.

Adult↗

Electrophysiologic correlates of direction and elevation cues for sound localization in the human brainstem.

Our objective was to study the effects of sound source direction and elevation on human brainstem electrical activity associated with sound localization. The subjects comprised 15 normal-hearing and symmetrically hearing adults Auditory brainstem evoked potentials (ABEPs) were recorded from three channels, in response to alternating-polarity clicks, presented at a rate of 21.1/s, at nine virtual spatial locations with different direction and elevation attributes Equivalent dipoles of the binaural interaction components (BICs) of ABEPs were derived by subtracting the response to binaural clicks at each spatial location from the algebraic sum of monaural responses to stimulation of each ear in turn. The BICs included two major components corresponding in latency to the vertex-neck-recorded components V and VI of ABEP. A significant decrease of the first BIC's equivalent dipole magnitude was observed for clicks in the horizontal-frontal position (no elevation) in the midsagittal plane, and for clicks in the left-horizontal (no elevation) and right diagonally above the head (medium elevation) positions in the coronal plane, compared to clicks positioned directly above the head. Significant effects on equivalent dipole latencies of this component were found for front-back positions in the midsagittal plane and left-right positions in the coronal plane, compared to clicks positioned directly above the head. The most remarkable finding was a significant change in equivalent dipole orientations across stimulus conditions. We conclude that the changes in BIC equivalent dipole latency, amplitude and orientation across stimulus conditions reflect differences in the distribution of binaural pontine activity evoked by sounds in different spatial locations.

Adult↗

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.

Acoustic Stimulation↗

Sound localization under perturbed binaural hearing.

This paper reports on the acute effects of a monaural plug on directional hearing in the horizontal (azimuth) and vertical (elevation) planes of human listeners. Sound localization behavior was tested with rapid head-orienting responses toward brief high-pass filtered (>3 kHz; HP) and broadband (0.5-20 kHz; BB) noises, with sound levels between 30 and 60 dB, A-weighted (dBA). To deny listeners any consistent azimuth-related head-shadow cues, stimuli were randomly interleaved. A plug immediately degraded azimuth performance, as evidenced by a sound level-dependent shift ("bias") of responses contralateral to the plug, and a level-dependent change in the slope of the stimulus-response relation ("gain"). Although the azimuth bias and gain were highly correlated, they could not be predicted from the plug's acoustic attenuation. Interestingly, listeners performed best for low-intensity stimuli at their normal-hearing side. These data demonstrate that listeners rely on monaural spectral cues for sound-source azimuth localization as soon as the binaural difference cues break down. Also the elevation response components were affected by the plug: elevation gain depended on both stimulus azimuth and on sound level and, as for azimuth, localization was best for low-intensity stimuli at the hearing side. Our results show that the neural computation of elevation incorporates a binaural weighting process that relies on the perceived, rather than the actual, sound-source azimuth. It is our conjecture that sound localization ensues from a weighting of all acoustic cues for both azimuth and elevation, in which the weights may be partially determined, and rapidly updated, by the reliability of the particular cue.

Acoustic Stimulation↗

The inability of squirrel monkeys to localize sound after unilateral ablation of auditory cortex.

The ability of squirrel monkeys to localize brief sounds was tested before and after unilateral ablation of auditory cortex. Results indicated that, after unilateral ablation of auditory cortex, monkeys could no longer localize sound accurately in the hemifield opposite to the side of lesion. These data demonstrate that contralaterally-directed functioning of auditory cortex occurs in primates.

Animals↗

Sound localization in hemispherectomized patients.

In order to precisely evaluate the consequences of cortical damage on free-field sound localization in humans, the present study examined response accuracy to auditory targets in three hemispherectomized patients and IQ-matched controls. Listeners reported sound location by pointing with their dominant hand to the apparent sound location in an anechoic chamber. Two conditions were tested: (i) localization of a fixed-sound source and (ii) localization of the beginning and the end of a simulated moving stimulus. In both conditions, the responses of the patients were less accurate than those of the controls in the hemifield contralateral to their removed hemisphere. Moreover, the single-case analyses revealed that the performances obtained with fixed sources were generally more precise than those obtained with moving sources. This result is discussed in terms of a differential involvement of cortical and subcortical pathways in the processing of stationary and moving sounds. Finally, the age at surgery and the post-surgical interval were related with the magnitude of the deficits, suggesting the possible influences of functional reorganization and cerebral plasticity.

Adolescent↗

Sound localization and sensitivity to interaural cues in bilateral users of the Med-El Combi 40/40+cochlear implant system.

OBJECTIVE: The purpose of this study was to investigate sound localization in subjects bilaterally implanted with MED-EL COMBI 40/40+ cochlear implants. In addition, the sensitivity to interaural cues was assessed. METHODS: In the localization test (11 subjects), CCITT noise (500 ms, original and HRTF-filtered, 70/75/80 dB sound pressure level) was presented from one of seven loudspeakers between -90 degrees and 90 degrees azimuth. The subject had to indicate which loudspeaker the noise was presented from. Sensitivity to interaural level differences (ILD) was assessed by performing localization tests (4 subjects) with the loudness of the two speech processors unbalanced to various degrees. To investigate the subjects' sensitivity to interaural time differences (ITD), lateralization was measured (7 subjects) as a function of the time difference between two Gaussian-like pulses, each directed to one of the subject's speech processor microphones by way of headphones. RESULTS: The judgments of all subjects significantly correlated with the positions of the loudspeakers. The scatter in the judged azimuth measured by the standard deviation of the responses was on average 27.5 degrees . Unbalanced loudness of the speech processors produced a bias in azimuth toward the speech processor with the louder volume setting. The mean rate of shift was 1.4 degrees per unit on the Wuerzburg loudness scale. Six of seven subjects showed a significant sensitivity to ITDs with the approximate time difference required for complete lateralization being 1,200 micros on an average. The one subject not showing a sensitivity to ITDs performed worst in the localization test. CONCLUSIONS: Bilateral cochlear implantation can restore spatial hearing in cochlear implant users. Both ILDs and ITDs are used by bilateral cochlear implant users in sound localization with ILDs appearing to be the dominant cue.

Adolescent↗

Cortical control of sound localization in the cat: unilateral cooling deactivation of 19 cerebral areas.

We examined the ability of mature cats to accurately orient to, and approach, an acoustic stimulus during unilateral reversible cooling deactivation of primary auditory cortex (AI) or 1 of 18 other cerebral loci. After attending to a central visual stimulus, the cats learned to orient to a 100-ms broad-band, white-noise stimulus emitted from a central speaker or 1 of 12 peripheral sites (at 15 degrees intervals) positioned along the horizontal plane. Twenty-eight cats had two to six cryoloops implanted over multiple cerebral loci. Within auditory cortex, unilateral deactivation of AI, the posterior auditory field (PAF) or the anterior ectosylvian sulcus (AES) resulted in orienting deficits throughout the contralateral field. However, unilateral deactivation of the anterior auditory field, the second auditory cortex, or the ventroposterior auditory field resulted in no deficits on the orienting task. In multisensory cortex, unilateral deactivation of neither ventral or dorsal posterior ectosylvian cortices nor anterior or posterior area 7 resulted in any deficits. No deficits were identified during unilateral cooling of the five visual regions flanking auditory or multisensory cortices: posterior or anterior ii suprasylvian sulcus, posterior suprasylvian sulcus or dorsal or ventral posterior suprasylvian gyrus. In motor cortex, we identified contralateral orienting deficits during unilateral cooling of lateral area 5 (5L) or medial area 6 (6m) but not medial area 5 or lateral area 6. In a control visual-orienting task, areas 5L and 6m also yielded deficits to visual stimuli presented in the contralateral field. Thus the sound-localization deficits identified during unilateral deactivation of area 5L or 6m were not unimodal and are most likely the result of motor rather than perceptual impairments. Overall, three regions in auditory cortex (AI, PAF, AES) are critical for accurate sound localization as assessed by orienting.

Acoustic Stimulation↗

Sound localization in the presence of one or two distracters.

Localizing a target sound can be a challenge when one or more distracter sounds are present at the same time. This study measured the effect of distracter position on target localization for one distracter (17 positions) and two distracters (21 combinations of 17 positions). Listeners were instructed to point to the apparent position of a train of 30-ms noise bursts, presented at 1 of 85 positions in virtual free field. A harmonic complex and a frequency-swept complex tone served as distracters. The two distracters were turned on 40 and 80 ms after the target onset, had temporal envelopes similar to that of the target, and did not overlap temporally with the target. Virtual sounds were synthesized with individual HRTFs. Localization performance degraded as the number of distracters increased from 0 to 2. When the horizontal distance between target and a single distracter was small (i.e., the interaural differences were almost the same), the influence on the apparent position was greater than when they were far apart. In the vertical dimension, there was not a systematic effect of distracter position on target localizability. However, there was a substantial increase in localization error for targets at high elevations (above 30 degrees) when distracters were present.

Attention↗

Neural substrates of sound localization.

Recent behavioral experiments suggest that parallel pathways in the forebrain and midbrain are capable of mediating sound localization. Neurophysiological studies have advanced our understanding of how binaural and monaural localization cues contribute to the representations of auditory space within these pathways. Experience-dependent modifications of auditory spatial tuning in both the forebrain and midbrain pathways indicate that sensory input during development customizes an individual's neural processing of spatial information.

Animals↗

Role of the dorsal cochlear nucleus in the sound localization behavior of cats.

The role of the dorsal cochlear nucleus (DCN) in directional hearing was evaluated by measuring sound localization behaviors before and after cats received lesions of the dorsal and intermediate acoustic striae (DAS/IAS). These lesions are presumed to disrupt spectral processing in the DCN without affecting binaural time and level difference cues that exit the cochlear nucleus via the ventral acoustic stria. Prior to DAS/IAS lesions, cats made accurate head orientation responses toward sound sources in the frontal sound field. After a unilateral DAS/IAS lesion, subjects showed increased errors in the azimuth and elevation of their responses; in addition, the final orientation of head movements tended to be more variable. Largest deficits in response elevation were observed in the hemifield that was ipsilateral to the lesion. When a second lesion was placed in the opposite DAS/IAS, increased orientation errors were observed throughout the frontal field. Nonetheless, bilaterally lesioned cats showed normal discrimination of changes in sound source location when tested with a spatial acuity task. These findings support previous interpretations that the DCN contributes to sound orientation behavior, and further suggest that the identification of absolute sound source locations and the discrimination between spatial locations involve independent auditory processing mechanisms.

Animals↗

Spectral cues for sound localization in cats: a model for discharge rate representations in the auditory nerve.

Neural representations of pinna-based spectral cues for sound localization were modeled by simulating auditory nerve discharge rates to noise bursts that had been shaped by filtering properties of the cat's head-related transfer functions (HRTFs) at 179 locations in the frontal field. The auditory nerve model transformed spectral differences between HRTFs into simulated neural rate differences. Linear equations for this transformation were developed from actual auditory nerve responses to a limited subset of HRTF-filtered noise bursts [Rice et al., J. Acoust. Soc. Am. 97, 1764-1776 (1995)]. Signal detection methods were used to investigate simulated neural responses to pairwise changes between HRTFs. The quality of neural representation for these changes, in terms of d' values, declined when the reference HRTF was moved from a central location (0 degree AZ, 0 degree EL) to a large positive azimuth in the horizontal plane (75 degrees AZ, 0 degree EL) or a high elevation in the median plane (0 degree AZ, 75 degrees EL). Most simulated responses exhibited large d' values for comparisons of contralateral versus ipsilateral azimuths, or eccentric versus frontal elevations. This rate information resulted from directionally dependent changes in the overall gain of HRTFs. In addition, fibers with best frequency (BF: the frequency of greatest sensitivity for individual fibers) between 5 and 18 kHz showed large d' values for HRTF contrasts in the immediate frontal field because of the effects of spectral notches (i.e., sharp drops in gain over a narrow frequency range). Spectral notches also played a prominent role in simulations that required identification of HRTF location in the absence of a fixed reference stimulus. These modeling results correspond well with previously described patterns in the cat's localization behaviors.

Animals↗

Discrimination of sound localization cues in young infants.

Infants 8 and 16 weeks old were tested in a visual fixation paradigm for the detection and use of sound localization cues. The effects of changes in both interaural intensity and interaural arrival time were assessed independently. For each trial, sounds were presented at the same time as visual displays located on both sides of the infant. The first eye movement and total looking time to each of the visual displays were noted by corneal reflection. Following a series of trials in which the apparent location of the sound was on 1 side, the apparent location was changed for a series of test trials. Neither age group responded with appropriate directional eye movements. The older infants did, however, show generally increased looking time on the test trials for both binaural cues. The 8-week-olds responded on test trials only when the interaural arrival time was changed. The data indicate that both binaural cues can be detected by 16-week-old infants although interaural arrival time was more salient in this paradigm.

Auditory Perception↗

Vertical-plane sound localization probed with ripple-spectrum noise.

Ripple-spectrum stimuli were used to investigate the scale of spectral detail used by listeners in interpreting spectral cues for vertical-plane localization. In three experiments, free-field localization judgments were obtained for 250-ms, 0.6-16-kHz noise bursts with log-ripple spectra that varied in ripple density, peak-to-trough depth, and phase. When ripple density was varied and depth was held constant at 40 dB, listeners' localization error rates increased most (relative to rates for flat-spectrum targets) for densities of 0.5-2 ripples/oct. When depth was varied and density was held constant at 1 ripple/oct, localization accuracy was degraded only for ripple depths > or = 20 dB. When phase was varied and density was held constant at 1 ripple/oct and depth at 40 dB, three of five listeners made errors at consistent locations unrelated to the ripple phase, whereas two listeners made errors at locations systematically modulated by ripple phase. Although the reported upper limit for ripple discrimination is 10 ripples/oct [Supin et al., J. Acoust. Soc. Am. 106, 2800-2804 (1999)], present results indicate that details finer than 2 ripples/oct or coarser than 0.5 ripples/oct do not strongly influence processing of spectral cues for sound localization. The low spectral-frequency limit suggests that broad-scale spectral variation is discounted, even though components at this scale are among those contributing the most to the shapes of directional transfer functions.

Acoustic Stimulation↗

Parallel pathways mediating both sound localization and gaze control in the forebrain and midbrain of the barn owl.

The hypothesis that sound localization and gaze control are mediated in parallel in the midbrain and forebrain was tested in the barn owl. The midbrain pathway for gaze control was interrupted by reversible inactivation (muscimol injection) or lesion of the optic tectum. Auditory input to the forebrain was disrupted by reversible inactivation or lesion of the primary thalamic auditory nucleus, nucleus ovoidalis (homolog of the medial geniculate nucleus). Barn owls were trained to orient their gaze toward auditory or visual stimuli presented from random locations in a darkened sound chamber. Auditory and visual test stimuli were brief so that the stimulus was over before the orienting response was completed. The accuracy and kinetics of the orienting responses were measured with a search coil attached to the head. Unilateral inactivation of the optic tectum had immediate and long-lasting effects on auditory orienting behavior. The owls failed to respond on a high percentage of trials when the auditory test stimulus was located on the side contralateral to the inactivated tectum. When they did respond, the response was usually (but not always) short of the target, and the latency of the response was abnormally long. When the auditory stimulus was located on the side ipsilateral to the inactivated tectum, responses were reliable and accurate, and the latency of responses was shorter than normal. In a tectally lesioned animal, response probability and latency to contralateral sounds returned to normal within 2 weeks, but the increase in response error (due to undershooting) persisted for at least 12 weeks. Despite abnormalities in the response, all of the owls were capable of localizing and orienting to contralateral auditory stimuli on some trials with the optic tectum inactivated or lesioned. This was not true for contralateral visual stimuli. Immediately following tectal inactivation, the owls exhibited complete neglect for visual stimuli located more than 20 degrees to the contralateral side (i.e., beyond the edge of the visual field of the ipsilateral eye). In the tectally lesioned animal, this neglect diminished with time. Unilateral inactivation of nucleus ovoidalis had different effects in three owls. Response error to contralateral sound sources increased for one owl and decreased for two; response error to ipsilateral sources did not change significantly for any. The probability of response to ipsilateral (but not contralateral) stimuli decreased for one owl. The latency of response to ipsilateral (but not contralateral) stimuli increased for one and decreased for another. All of the owls, however, routinely localized and oriented toward ipsilateral and contralateral auditory stimuli with nucleus ovoidalis inactivated.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

Influences of age and hearing loss on the precedence effect in sound localization.

Cranford, Boose, & Moore (1990a) reported that many elderly persons exhibit problems in perceiving the apparent location of fused auditory images in a sound localization task involving the Precedence Effect (PE). In the earlier study, differences in peripheral hearing sensitivity between young and elderly subjects were not controlled. In the present study, four groups of young and elderly subjects, matched with respect to age and the presence or absence of sensorineural hearing loss, were examined to determine the effects of these two factors on performance with the PE task. Although significantly poorer performances on the PE task were found to be associated with both increased age and hearing loss, additional tentative evidence was obtained that the presence of hearing loss may have a relatively greater detrimental effect on the performance of at least some elderly subjects than it does on younger persons.

Adult↗

Auditory processing of spectral cues for sound localization in the inferior colliculus.

The head-related transfer function (HRTF) of the cat adds directionally dependent energy minima to the amplitude spectrum of complex sounds. These spectral notches are a principal cue for the localization of sound source elevation. Physiological evidence suggests that the dorsal cochlear nucleus (DCN) plays a critical role in the brainstem processing of this directional feature. Type O units in the central nucleus of the inferior colliculus (ICC) are a primary target of ascending DCN projections and, therefore, may represent midbrain specializations for the auditory processing of spectral cues for sound localization. Behavioral studies confirm a loss of sound orientation accuracy when DCN projections to the inferior colliculus are surgically lesioned. This study used simple analogs of HRTF notches to characterize single-unit response patterns in the ICC of decerebrate cats that may contribute to the directional sensitivity of the brain's spectral processing pathways. Manipulations of notch frequency and bandwidth demonstrated frequency-specific excitatory responses that have the capacity to encode HRTF-based cues for sound source location. These response patterns were limited to type O units in the ICC and have not been observed for the projection neurons of the DCN. The unique spectral integration properties of type O units suggest that DCN influences are transformed into a more selective representation of sound source location by a local convergence of wideband excitatory and frequency-tuned inhibitory inputs.

Acoustic Stimulation↗