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Auditory localization behaviour in visually deprived cats.

The ability to localize sounds in azimuth was tested in five cats that had been binocularly deprived of vision from birth for several months and in three normal age-matched controls. Brief tone bursts were presented in an eight-choice apparatus along 360 degrees of the azimuthal plane at constant elevation. Using positive reinforcement techniques, the cats were trained to walk from the centre of the 3 m diameter circular enclosure to the hidden loudspeakers. The distribution of sound localization error from 55 trials per cat at each speaker position was measured, and its standard deviation was used to assess the precision of sound localization. All cats localized tones straight ahead of them most precisely; performance at lateral and rear positions was gradually less precise. When the sound localization ability of normal and binocularly deprived cats was compared across speakers, a significantly enhanced precision was found for binocularly deprived cats overall (P < 0.002; two-way analysis of variance). An improvement was found at each individual speaker position, but it was greatest at lateral and rear positions. In two sets of control experiments normal cats were retested (i) in the dark with the aid of an infrared camera and (ii) after 3 months of binocular lid suture. Normal cats in the dark did not show any differences in their sound localization behaviour. Late-deprived cats showed a tendency for better performance, which fell short of statistical significance. Our results in visually deprived cats agree well with some reports on the sound localization ability of blind humans, but disagree with others. Our data provide support for a hypothesis of compensatory plasticity, in which sensory functions get sharpened with the loss of another modality. They seem to rule out the necessity for vision to play a role in the postnatal calibration of auditory space.

Acoustic Stimulation↗

Identification and localization of sound sources in the median sagittal plane.

The ability of human listeners to identify broadband noises differing in spectral structure was studied for multiple sound-source locations in the median sagittal plane. The purpose of the study was to understand how sound identification is affected by spectral variations caused by directionally dependent head-related transfer functions. It was found that listeners could accurately identify noises with different spectral peaks and valleys when the source location was fixed. Listeners could also identify noises when the source location was roved in the median sagittal plane when the relevant spectral features were at low frequency. Listeners failed to identify noises with roved location when the spectral structure was at high frequency, presumably because the spectral structure was confused with the spectral variations caused by different locations. Parallel experiments on sound localization showed that listeners can localize noises that they cannot identify. The combination of identification and localization experiments leads to the conclusion that listeners cannot compensate for directionally dependent filtering by their own heads when they try to identify sounds.

Adult↗

Localizing nearby sound sources in a classroom: binaural room impulse responses.

Binaural room impulse responses (BRIRs) were measured in a classroom for sources at different azimuths and distances (up to 1 m) relative to a manikin located in four positions in a classroom. When the listener is far from all walls, reverberant energy distorts signal magnitude and phase independently at each frequency, altering monaural spectral cues, interaural phase differences, and interaural level differences. For the tested conditions, systematic distortion (comb-filtering) from an early intense reflection is only evident when a listener is very close to a wall, and then only in the ear facing the wall. Especially for a nearby source, interaural cues grow less reliable with increasing source laterality and monaural spectral cues are less reliable in the ear farther from the sound source. Reverberation reduces the magnitude of interaural level differences at all frequencies; however, the direct-sound interaural time difference can still be recovered from the BRIRs measured in these experiments. Results suggest that bias and variability in sound localization behavior may vary systematically with listener location in a room as well as source location relative to the listener, even for nearby sources where there is relatively little reverberant energy.

Acoustics↗

Directional hearing in aging rats.

The loss of the ability to localize sounds was discovered to occur in rats as a consequence of aging. Subjects were trained in a two-alternative forced-choice paradigm to press the left bar if a sound was presented from the left side and to press the right bar if sound was presented from the right side. Testing was conducted with a brief 200 msec pink noise signal. The accuracy of sound localization was measured in rats from 10 to 21 months of age. By 21 months of age the accuracy of localization had declined from greater than 90% correct to less than 69% correct. The loss of the ability to localize sounds may be involved with the onset of spatial disorientation observed in some subjects at advanced ages.

Aging↗

Asymmetric performances in binaural localization of sound in space.

Twenty right-handers and 20 left-handers were tested on a sound localization task. Broadband noise was presented from either the left or right hemifield. Localization accuracy was significantly greater (P = 0.002) when sounds emanated from the left hemifield thereby suggesting a paramount role played by the right hemisphere. Correcting for front-rear reversals, attributable to impoverished spectral cues and/or faulty processing of such cues, rendered differences in error scores linked to hemifield nonsignificant. The data were interpreted to mean that the special contribution of the right hemisphere to this task was its greater fidelity in processing spectral cues. No differences in localization proficiency between right- and left-handers were observed.

Adult↗

Sound source localization in real sound fields based on empirical statistics of interaural parameters.

The role of temporal fluctuations and systematic variations of interaural parameters in localization of sound sources in spatially distributed, nonstationary noise conditions was investigated. For this, Bayesian estimation was applied to interaural parameters calculated with physiologically plausible time and frequency resolution. Probability density functions (PDFs) of the interaural level differences (ILDs) and phase differences (IPDs) were estimated by measuring histograms for a directional sound source perturbed by several types of interfering noise at signal-to-noise ratios (SNRs) between -5 and +30 dB. A moment analysis of the PDFs reveals that the expected values shift and the standard deviations increase considerably with decreasing SNR, and that the PDFs have non-Gaussian shape at medium SNRs. A d' analysis of the PDFs indicates that elevation discrimination is possible even at low SNRs in the median plane by integrating information across frequency. Absolute sound localization was simulated by a Bayesian maximum a posteriori (MAP) procedure. The simulation is based on frequency integration of broadly tuned "detectors." Confusion patterns of real and estimated sound source directions are similar to those of human listeners. The results indicate that robust processing strategies are needed to exploit interaural parameters successfully in noise conditions due to their strong temporal fluctuations.

Acoustic Stimulation↗

Localization by unilateral BAHA users.

OBJECTIVES: Patients with unilateral hearing loss report difficulty hearing conversation on their impaired side, localizing sound, and understanding of speech in background noise. The bone-anchored cochlear stimulator (BAHA) (Entific, Gothenburg, Sweden) has been shown to improve performance in persons with unilateral severe-profound sensorineural loss (USNHL). The purpose of this study is to evaluate the effectiveness of BAHA in sound localization for USNHL listeners. STUDY DESIGN: Prospective study of 12 USNHL subjects, 9 of whom received implants on the poorer hearing side. A control group of 10 normal hearing subjects were assessed for comparison. Localization with and without BAHA was assessed using an array of 8 speakers at head level separated by 45 degrees. Error analysis matrix was generated to evaluate the confusions, accuracy in response, and laterality judgment. RESULTS: The average accuracy of speaker localization was 16% in the unaided condition, with no improvement with BAHA use. Laterality judgment was poorer than 43% in both aided and nonaided conditions. CONCLUSIONS: Patients with UNSNHL had poor sound localization and laterality judgment abilities that did not improve with BAHA use.

Aged↗

Effect of eye position on saccades and neuronal responses to acoustic stimuli in the superior colliculus of the behaving cat.

We examined the motor error hypothesis of visual and auditory interaction in the superior colliculus (SC), first tested by Jay and Sparks in the monkey. We trained cats to direct their eyes to the location of acoustic sources and studied the effects of eye position on both the ability of cats to localize sounds and the auditory responses of SC neurons with the head restrained. Sound localization accuracy was generally not affected by initial eye position, i.e., accuracy was not proportionally affected by the deviation of the eyes from the primary position at the time of stimulus presentation, showing that eye position is taken into account when orienting to acoustic targets. The responses of most single SC neurons to acoustic stimuli in the intact cat were modulated by eye position in the direction consistent with the predictions of the "motor error" hypothesis, but the shift accounted for only two-thirds of the initial deviation of the eyes. However, when the average horizontal sound localization error, which was approximately 35% of the target amplitude, was taken into account, the magnitude of the horizontal shifts in the SC auditory receptive fields matched the observed behavior. The modulation by eye position was not due to concomitant movements of the external ears, as confirmed by recordings carried out after immobilizing the pinnae of one cat. However, the pattern of modulation after pinnae immobilization was inconsistent with the observations in the intact cat, suggesting that, in the intact animal, information about the position of the pinnae may be taken into account.

Acoustic Stimulation↗

A spatial hearing deficit in early-blind humans.

An important issue in neuroscience is the effect of visual loss on the remaining senses. Two opposing views have been advanced. On the one hand, visual loss may lead to compensatory plasticity and sharpening of the remaining senses. On the other hand, early blindness may also prevent remaining sensory modalities from a full development. In the case of sound localization, it has been reported recently that, under certain conditions, early-blind humans can localize sounds better than sighted controls. However, these studies were confined to a single sound source in the horizontal plane. This study compares sound localization of early-blind and sighted subjects in both the horizontal and vertical domain, whereas background noise was added to test more complex hearing conditions. The data show that for high signal-to-noise (S/N) ratios, localization by blind and sighted subjects is similar for both azimuth and elevation. At decreasing S/N ratios, the accuracy of the elevation response components deteriorated earlier than the accuracy of the azimuth component in both subject groups. However, although azimuth performance was identical for the two groups, elevation accuracy deteriorated much earlier in the blind subject group. These results indicate that auditory hypercompensation in early-blind humans does not extend to the frontal target domain, where the potential benefit of vision is maximal. Moreover, the results demonstrate for the first time that in this domain the human auditory system may require vision to optimally calibrate the elevation-related spectral pinna cues. Sensitivity to azimuth-encoding binaural difference cues, however, may be adequately calibrated in the absence of vision.

Acoustic Stimulation↗

The functional organization of auditory working memory as revealed by fMRI.

Spatial and nonspatial auditory tasks preferentially recruit dorsal and ventral brain areas, respectively. However, the extent to which these auditory differences reflect specific aspects of mental processing has not been directly studied. In the present functional magnetic resonance imaging experiment, participants encoded and maintained either the location or the identity of a sound for a delay period of several seconds and then subsequently compared that information with a second sound. Relative to sound localization, sound identification was associated with greater hemodynamic activity in the left rostral superior temporal gyrus. In contrast, localizing sounds recruited greater activity in the parietal cortex, posterior temporal lobe, and superior frontal sulcus. The identification differences were most prominent during the early stage of the trial, whereas the location differences were most evident during the late (i.e., comparison) stage. Accordingly, our results suggest that auditory spatial and identity dissociations as revealed by functional imaging may be dependent to some degree on the type of processing being carried out. In addition, dorsolateral prefrontal and lateral superior parietal areas showed greater activity during the comparison as opposed to the earlier stage of the trial, regardless of the type of auditory task, consistent with results from visual working memory studies.

Adult↗

Synaptic depression in the localization of sound.

Short-term synaptic plasticity, which is common in the central nervous system, may contribute to the signal processing functions of both temporal integration and coincidence detection. For temporal integrators, whose output firng rate depends on a running average of recent synaptic inputs, plasticity modulates input synaptic strength and thus may directly control signalling gain and the function of neural networks. But the firing probability of an ideal coincidence detector would depend on the temporal coincidence of events rather than on the average frequency of synaptic events. Here we have examined a specific case of how synaptic plasticity can affect temporal coincidence detection, by experimentally characterizing synaptic depression at the synapse between neurons in the nucleus magnocellularis and coincidence detection neurons in the nucleus laminaris in the chick auditory brainstem. We combine an empirical description of this depression with a biophysical model of signalling in the nucleus laminaris. The resulting model predicts that synaptic depression provides an adaptive mechanism for preserving interaural time-delay information (a proxy for the location of sound in space) despite the confounding effects of sound-intensity-related information. This mechanism may help nucleus laminaris neurons to pass specific sound localization information to higher processing centres.

Acoustic Stimulation↗

The role of phase changes in sound signals in localization of sound sources.

The auditory system in humans and animals makes virtually no discrimination of phase changes in the structure of monaurally presented sound signals. However, electrophysiological studies have demonstrated marked changes in the responses of the central parts of the auditory system when the phase structure of the signal changes during presentation of the same type of stimulation. We have suggested that this inconsistency is due to the preparative role of phase effects during monaural stimulation for subsequent operations in the auditory system involved in determining the location of a sound source in space. This report presents experimental data on defined changes (increases in amplitude) in the electrical responses of the midbrain center of the auditory system (inferior colliculus) in antiphase binaural presentation of series of sound impulses (comparison with synphase presentation). These changes may be part of the mechanism underlying the interference resistance of the auditory system during determination of the location of a sound source (binaural release from masking). Neuronal cortical activity is sensitive and selective to dynamic interaural changes in the phase spectrum of the signal, which may provide the basis of the mechanism for locating a moving sound source. Auditory evoked potentials in humans demonstrate memorizing of the direction of movement of a sound image, as shown by the changes in parameters on presentation of stimuli of different locations (deviant stimuli) differing from the standard parameters of mismatch negativity.

Acoustic Stimulation↗

A PET study of human auditory spatial processing.

To learn more about human auditory spatial processing, we used positron emission tomography (PET) to measure regional cerebral blood flow in human volunteers engaged in sound localization tasks. Spectral and binaural cues of localized sound were reproduced by a sound system and delivered via headphones. During localization tasks, subjects activated inferior parietal lobules (IPL) bilaterally. In a second experiment, matched in design to the first, subjects made non-spatial auditory discriminations based on frequency, activating the IPL bilaterally with left hemispheric predominance. A between-study comparison revealed that the right IPL was significantly more activated during the sound localization task compared with the feature discrimination task, suggesting a preferential role for the right IPL in auditory spatial processing.

Adult↗

Active localization of virtual sounds.

A simple virtual sound display built around a microcomputer and analog hardware is described. The display implements most of the primary cues for sound localization in the ear-level plane. Judging both from informal observations by users and from objective data obtained in an experiment on homing to virtual and real sounds, it is concluded that simple displays like the one described are effective in creating the impression of external sounds to which observers can locomote with ease; in particular, this means that simulation of the direction-dependent spectral shaping effects of the pinnae is not a necessary requirement for extracranial sound localization.

Adult↗

Cutaneous senses for detection and localization of environmental sound sources: a review and tutorial.

A review and tutorial on the potential use of the skin for sound localization are presented. Some basic physical and psychophysical properties are reviewed; special properties and phantom perceptions associated with cutaneous stimulation are presented; experiments by Békésy, Gescheider, Frost, Richardson and Weisenberger on different instrumentation for/and results on localization of sound sources with skin stimulation are analysed. The best results have been obtained using intensity differences between two vibrators. Sound source localization precision approaching that of the auditory sense in quiet surroundings has been obtained, particularly when free head movements are allowed. Significant results have also been obtained with temporal differences, provided the natural sound delay between the ears is increased to fit the coding characteristics of the skin. As yet, no portable equipment has been designed and tested in realistic environments.

Auditory Perception↗

Responses of single neurons in physiologically defined area AI of cat cerebral cortex: sensitivity to interaural intensity differences.

In 15 cats, cortical area AI was defined by its frequency organization, and cells within that field were tested for sensitivity to interaural intensity differences (IIDs) using sealed stimulus delivery systems. Of 39 cells tested quantitatively, 26 were sensitive to IIDs. In 70% of cases, sensitivity to IIDs reflected suppressive binaural interactions, and was manifested as a sigmoidal relation of spike count to IID. For 8 other cells, facilitative binaural interactions generated unit sensitivity to IIDS; three of these neurons demonstrated nonmonotonic dependency of spike count on IID, with peak firing rates at or near 0 dB IID. Analysis of spike count versus IID functions in terms of the auditory azimuths known to generate the IIDs used revealed that the majority of cells were most sensitive to IIDs associated with azimuths in the contralateral sound field. These data are compatible with other evidence on the sensitivity of cortical and brainstem cells to binaural sound localization cues, and suggest that each side of the auditory brain is independently capable of localizing sound sources in the contralateral field.

Animals↗

Vestigial hearing in a fossorial mammal, the pocket gopher (Geomys bursarius).

Behavioral tests of hearing and sound localization in the North American pocket gopher (Geomys bursarius) show that it is unique among mammals. It has a severely attenuated range of hearing and only rudimentary ability to localize sound. In these respects, the hearing of gophers can be properly termed 'vestigial' and suggests that life underground can produce as severe a change in hearing as a light-less world produces in vision or an odorless world produces in olfaction.

Acoustic Stimulation↗

Binaural processing after corrected congenital unilateral conductive hearing loss.

Binaural processing was measured in a series of tests in patients before and after surgery to correct congenital unilateral conductive hearing losses. Data are presented from 19 patients between the ages of 6 and 33 years that had an abnormal external and/or middle ear on one side but normal hearing in the other ear. Surgery improved thresholds an average of 36 dB HL (from 56 to 20 dB HL). Patients were tested pre- and postoperatively for interaural temporal difference limens, alternate and simultaneous loudness balances, sound localization, binaural detection thresholds, and speech perception in noise. There was statistically significant improvement after surgery in all tests, and the amount of improvement varied along a continuum that appears to be related to the simplicity of the task. For example, most postoperative patients had normal or near-normal performance in a test of interaural temporal difference limens, while almost all had difficulty localizing sounds. Neither binaural performance (before or after surgery) nor the improvement in performance was correlated with age, pure-tone thresholds, or asymmetry. Limited available data show no significant changes in performance from four weeks to over 24 weeks after surgery. In conclusion, binaural ability following corrective surgery exists in varying degrees in these tasks, suggesting different effects of abnormal early experience on different aspects of binaural hearing.

Adolescent↗