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PubMed2002

Contribution of spectral cues to human sound localization.

The contribution of spectral cues to human sound localization was investigated by removing cues in 1/2-, 1- or 2-octave bands in the frequency range above 4 kHz. Localization responses were given by placing an acoustic pointer at the same apparent position as a virtual target. The pointer was generated by filtering a 100-ms harmonic complex with equalized head-related transfer functions (HRTFs). Listeners controlled the pointer via a hand-held stick that rotated about a fixed point. In the baseline condition, the target, a 200-ms noise burst, was filtered with the same HRTFs as the pointer. In other conditions, the spectral information within a certain frequency band was removed by replacing the directional transfer function within this band with the average transfer of this band. Analysis of the data showed that removing cues in 1/2-octave bands did not affect localization, whereas for the 2-octave band correct localization was virtually impossible. The results obtained for the 1-octave bands indicate that up-down cues are located mainly in the 6-12-kHz band, and front-back cues in the 8-16-kHz band. The interindividual spread in response patterns suggests that different listeners use different localization cues. The response patterns in the median plane can be predicted using a model based on spectral comparison of directional transfer functions for target and response directions.

Adult↗
PubMed2003

Electrophysiological correlates of azimuth and elevation cues for sound localization in human middle latency auditory evoked potentials.

OBJECTIVE: To study, in humans, the effects of sound source azimuth and elevation on primary auditory cortex binaural activity associated with sound localization. DESIGN: Middle Latency Auditory Evoked Potentials (MLAEPs) were recorded from three channels, in response to alternating polarity clicks, presented at a rate of 5/sec, at nine virtual spatial locations with different azimuths and elevations. Equivalent dipoles of Binaural Interaction Components (BICs) of MLAEPs were derived from 15 normally and symmetrically hearing adults by subtracting the response to binaural clicks at each spatial location from the algebraic sum of responses to stimulation of each ear alone. The amplified potentials were averaged over 4000 repetitions using a dwell time of 78 micro sec/address/channel. Variations in magnitudes, latencies and orientations of the dipole equivalents of cortical activity were noted in response to the nine spatial locations. RESULTS: Middle-latency BICs included six major components corresponding in latency to the vertex-neck recorded components of MLAEP. A significant decrease of equivalent dipole magnitude was observed for two of the components: Pa2 in response to clicks in the backward positions (medium and no elevation); and Nb in response to clicks in the back and front positions (medium and no elevation) in the midsagittal plane. In the coronal plane, Pa2 equivalent dipole magnitude significantly decreased in response to right-horizontal (no elevation) clicks. Significant effects on equivalent dipole latencies of Pa2 were found for backward positions (no elevation) in the midsagittal plane. No significant effects on Pa2 and Nb equivalent dipole orientations were found across stimulus conditions. CONCLUSIONS: The changes in equivalent dipole magnitudes and latencies of MLAEP BICs across stimulus conditions may reflect spectral tuning in binaural primary auditory cortex neurons processing the frequency cues for sound localization.

Adult↗
PubMed2001

A neural code for low-frequency sound localization in mammals.

We report a systematic relationship between sound-frequency tuning and sensitivity to interaural time delays for neurons in the midbrain nucleus of the inferior colliculus; neurons with relatively low best frequencies (BFs) showed response peaks at long delays, whereas neurons with relatively high BFs showed response peaks at short delays. The consequence of this relationship is that the steepest region of the function relating discharge rate to interaural time delay (ITD) fell close to midline for all neurons irrespective of BF. These data provide support for a processing of the output of coincidence detectors subserving low-frequency sound localization in which the location of a sound source is determined by the activity in two broad, hemispheric spatial channels, rather than numerous channels tuned to discrete spatial positions.

Action Potentials↗
PubMed1984

A critical period for the recovery of sound localization accuracy following monaural occlusion in the barn owl.

We studied the ability of barn owls to recover accurate sound localization after being raised with one ear occluded. Most of the owls had ear plugs inserted before they reached adult size, and therefore they never experienced normal adult localization cues until their ear plugs were removed. Upon removal of their ear plugs, these owls exhibited large systematic sound localization errors. The rate at which they recovered accurate localization decreased with the age of the bird at the time of plug removal, and recovery essentially ceased when owls reached 38 to 42 weeks of age. We interpret this age as the end of a critical period for the consolidation of associations between auditory cues and locations in space. Owls that had experienced adult localization cues for a short period of time before ear plugging recovered normal accuracy rapidly, even if they remained plugged well past the end of the critical period. This suggests that a brief exposure to normal adult cues early in the critical period is sufficient to enable the recovery of localization accuracy much later in life.

Aging↗
PubMed1978

Sound localization: value in localizing lesions of the auditory pathway.

Reports in the literature suggest that patients with otosclerosis, neural lesions such as acoustic neuroma, pontine lesions, and temporal lobe lesions may show abnormal sound localization ability. In the present experiment, 15 normal subjects and 22 patients with various auditory pathway lesions had sound localization testing using special apparatus. The results confirmed that patients with otosclerosis were unable to localize low frequency sounds. Those with temporal lobe lesions made judgments not significantly different from normal subjects. Patients with unilateral sensorineural deafness made errors localizing high frequency sounds. Those with acoustic neuromas made greater errors than those without, but the difference between these two groups just failed to reach statistical significance. The number of cases was however small, and further studies are in progress to attempt to define numerical criteria to assist clinical decisions with regard to the selection of patients for invasive investigations such as oil cisternogram.

Deafness↗
PubMed1975

Sound localization by the bottlenose porpoise Tursiops truncatus.

1. Sound localization was measured behaviourally for the Atlantic bottlenose porpoise (Tursiops truncatus) using a wide range of pure tone pulses as well as clicks simulating the species echolocation click. 2. Measurements of the minimum audible angle (MAA) on the horizontal plane give localization discrimination thresholds of between 2 and 3 degrees for sounds from 20 to 90 kHz and thresholds from 2-8 to 4 degrees at 6, 10 and 100 kHz. With the azimuth of the animal changed relative to the speakers the MAAs were 1-3-1-5 degrees at an azimuth of 15 degrees and about 5 degrees for an azimuth of 30 degrees. 3. MAAs to clicks were 0-7-0-8 degrees. 4. The animal was able to do almost as well in determining the position of vertical sound sources as it could for horizontal localization. 5. The data indicate that at low frequencies the animal may have been localizing by using the region around the external auditory meatus as a detector, but at frequencies about 20 kHz it is likely that the animal was detecting sounds through the lateral sides of the lower jaw. 6. Above 20 kHz, it is likely that the animal was localizing using binaural intensity cues. 7. Our data support evidence that the lower jaw is an important channel for sound detection in Tursiops.

Animals↗
PubMed1977

Effects of unilateral ablation of auditory cortex on monaural cat's ability to localize sound.

1. Cats with one cochlea destroyed were trained to localize sound. After behavioral measures of the animal's accuracy of localization were made, cortical auditory areas were ablated unilaterally. 2. The results showed: a) like binaural localization, monaural localization of sound in space, as measured by the ability of an animal to move toward a sound source, depends on integrity of auditory cortex; b) it is only ablation of cortex contralateral to the functional ear that seriously affects localizing behavior; ablation of cortex ipsilateral to the intact cochlea has little or no effect on localizing behavior. 3. To explain the results, we suggest that auditory cortex is essential for an organized perception of space including the relation of the animal's position to other objects in space. We also suggest that auditory cortex contralateral to a given ear is necessary in order for the animal to recognize that a stimulus is presented to that ear of, when both ears are intact, to recognize that the stimulus to the given ear differs in some way (intensity, time of arrival, sequential arrangement of sounds) from the stimulus to the opposite ear.

Animals↗
PubMed1993

Sound localization in acallosal human listeners.

In order to evaluate the callosal and hemispheric involvement in sound localization, the present study examined response accuracy to auditory targets in acallosal subjects. The primary interest was to determine whether the congenital absence of the corpus callosum affects auditory localization, especially for sounds situated near the midline of auditory space or moving across it. A corollary objective was to examine the possible existence of an hemispheric asymmetry on audio-spatial localization tasks. Four subjects with callosal agenesis paired to four age and IQ-matched controls and 16 normal control subjects were asked to locate broad band noise bursts at fixed intensity (52 dB sound pressure level) in the horizontal plane in an anechoic chamber. Broad band noise bursts were delivered randomly through 16 loudspeakers, which were mounted at approximately 10 degrees intervals on a perimeter frame. Two conditions were tested: (i) localization of a fixed-sound source; (ii) localization of the beginning and the end of a simulated moving stimulus. Two response modes were used. Listeners reported the apparent stimulus location either (i) by pointing with the ipsilateral index finger or (ii) by calling out the estimated angles indicated on the calibrated sound perimeter. Aiming accuracy was assessed by calculating the mean deviation of the response from the objective target position. The results indicated that the responses of the acallosal subjects were less accurate than those of the controls. The deficit was observed not only at the midline but throughout the auditory field. This points to possible compensatory mechanisms following the early absence of the corpus callosum which are, however, limited. The results obtained with manual pointing were generally more precise than those obtained through oral responses. This difference suggests that the remapping of spatial positions onto a verbally based coordinate system involves a supplementary cognitive step which affects the precision of the response. Comparing the performance to stimulus presentation in the left and right fields indicated that no hemispheric asymmetry was apparent under any of the conditions for either the acallosal subjects or the IQ-matched and normal control subjects.

Adolescent↗
PubMed2006

A probabilistic model for binaural sound localization.

This paper proposes a biologically inspired and technically implemented sound localization system to robustly estimate the position of a sound source in the frontal azimuthal half-plane. For localization, binaural cues are extracted using cochleagrams generated by a cochlear model that serve as input to the system. The basic idea of the model is to separately measure interaural time differences and interaural level differences for a number of frequencies and process these measurements as a whole. This leads to two-dimensional frequency versus time-delay representations of binaural cues, so-called activity maps. A probabilistic evaluation is presented to estimate the position of a sound source over time based on these activity maps. Learned reference maps for different azimuthal positions are integrated into the computation to gain time-dependent discrete conditional probabilities. At every timestep these probabilities are combined over frequencies and binaural cues to estimate the sound source position. In addition, they are propagated over time to improve position estimation. This leads to a system that is able to localize audible signals, for example human speech signals, even in reverberating environments.

Artificial Intelligence↗
PubMed2006

Sound localization by barn owls in a simulated echoic environment.

We examined the accuracy and precision with which the barn owl (Tyto alba) turns its head toward sound sources under conditions that evoke the precedence effect (PE) in humans. Stimuli consisted of 25-ms noise bursts emitted from two sources, separated horizontally by 40 degrees, and temporally by 3-50 ms. At delays from 3 to 10 ms, head turns were always directed at the leading source, and were nearly as accurate and precise as turns toward single sources, indicating that the leading source dominates perception. This lead dominance is particularly remarkable, first, because on some trials, the lagging source was significantly higher in amplitude than the lead, arising from the directionality of the owl's ears, and second, because the temporal overlap of the two sounds can degrade the binaural cues with which the owl localizes sounds. With increasing delays, the influence of the lagging source became apparent as the head saccades became increasingly biased toward the lagging source. Furthermore, on some of the trials at delays > or = 20 ms, the owl turned its head, first, in the direction of one source, and then the other, suggesting that it was able to resolve two separately localizable sources. At all delays <50 ms, response latencies were longer for paired sources than for single sources. With the possible exception of response latency, these findings demonstrate that the owl exhibits precedence phenomena in sound localization similar to those in humans and cats, and provide a basis for comparison with neurophysiological data.

Acoustic Stimulation↗
PubMed2000

Correlation between the activity of single auditory cortical neurons and sound-localization behavior in the macaque monkey.

Lesion studies have indicated that the auditory cortex is crucial for the perception of acoustic space, yet it remains unclear how these neurons participate in this perception. To investigate this, we studied the responses of single neurons in the primary auditory cortex (AI) and the caudomedial field (CM) of two monkeys while they performed a sound-localization task. Regression analysis indicated that the responses of approximately 80% of neurons in both cortical areas were significantly correlated with the azimuth or elevation of the stimulus, or both, which we term "spatially sensitive." The proportion of spatially sensitive neurons was greater for stimulus azimuth compared with stimulus elevation, and elevation sensitivity was primarily restricted to neurons that were tested using stimuli that the monkeys also could localize in elevation. Most neurons responded best to contralateral speaker locations, but we also encountered neurons that responded best to ipsilateral locations and neurons that had their greatest responses restricted to a circumscribed region within the central 60 degrees of frontal space. Comparing the spatially sensitive neurons with those that were not spatially sensitive indicated that these two populations could not be distinguished based on either the firing rate, the rate/level functions, or on their topographic location within AI. Direct comparisons between the responses of individual neurons and the behaviorally measured sound-localization ability indicated that proportionally more neurons in CM had spatial sensitivity that was consistent with the behavioral performance compared with AI neurons. Pooling the responses across neurons strengthened the relationship between the neuronal and psychophysical data and indicated that the responses pooled across relatively few CM neurons contain enough information to account for sound-localization ability. These data support the hypothesis that auditory space is processed in a serial manner from AI to CM in the primate cerebral cortex.

Action Potentials↗
PubMed1992

Narrow-band sound localization related to external ear acoustics.

Human subjects localized brief 1/6-oct bandpassed noise bursts that were centered at 6, 8, 10, and 12 kHz. All testing was done under binaural conditions. The horizontal component of subjects' responses was accurate, comparable to that for broadband localization, but the vertical and front/back components exhibited systematic errors. Specifically, responses tended to cluster within restricted ranges that were specific for each center frequency. The directional transfer functions of the subjects' external ears were measured for 360 horizontal and vertical locations. The spectra of the sounds that were present in the subjects' ear canals, the "proximal stimulus" spectra, were computed by combining the spectra of the narrow-band sound sources with the directional transfer functions for particular stimulus locations. Subjects consistently localized sounds to regions within which the associated directional transfer function correlated most closely with the proximal stimulus spectrum. A quantitative model was constructed that successfully predicted subjects' responses based on interaural level difference and spectral cues. A test of the model, using techniques adapted from signal detection theory, indicated that subjects tend to use interaural level difference and spectral shape cues independently, limited only by a slight spatial correlation of the two cues. A testing procedure is described that provides a quantitative comparison of various predictive models of sound localization.

Acoustics↗
PubMed1982

[Effects of head movements on sound localization in pseudo-phonically reversed hearing].

Measurements of sound localization in the lateral dimension were taken with or without head movements when the observer wore the electronic pseudophone in an anechoic chamber. Tones of various frequencies as stimuli were presented from the loudspeakers located on the horizontal plane of the observer's head and on the circle, 2 m in diameter, around the observer, who pointed with his hand the position from which he thought the sound originated. The results of three subjects showed that: when the inter-aural axis of the pseudophone was 180 degrees reversed, real and apparent locations of the sound were symmetrical with respect to the median of the observer's head under the head-fixed condition, but were symmetrical with respect to the position of the observer's head as a center under the head-free condition. The implication of the results obtained was discussed.

Adult↗
PubMed1991

Coding of interaural time differences of transients in auditory cortex of Rattus norvegicus: implications for the evolution of mammalian sound localization.

We obtained quantitative evidence on the coding of interaural time differences (ITDs) of click stimuli by 40 single neurons in the auditory cortex of anesthetized albino rats. Most of the neurons (31/40) received an excitatory input from the contralateral ear, and an inhibitory input from the ipsilateral ear (EI cells). These neurons expressed their sensitivity to ITDs in a sigmoidal relation between spike count and ITD, with maximal responses associated with contralateral-leading ITDs. The mean ITD dynamic range was 590 microseconds. The dynamic ranges typically encompassed at least part of the behaviorally-relevant range (about +/- 130 microseconds). Variations in ITD from 130 microseconds favoring one ear to 130 microseconds favoring the other ear caused spike response rate changes, on average, of 29.5%. These data are similar to those previously presented for the central auditory systems of larger mammals, whose auditory localization acuity is significantly better than that of the rat. We argue, therefore, that the sound localization mechanisms based on transient ITDs have not evolved in a fashion that covaries with interaural distance, and that there exists a mismatch between the ITDs the rat will encounter in the free field, and the ITDs which are encoded by its nervous system. This may be one reason why sound localization acuity has a roughly inverse relation to interaural distance.

Acoustic Stimulation↗
PubMed2004

Dynamic sound localization during rapid eye-head gaze shifts.

Human sound localization relies on implicit head-centered acoustic cues. However, to create a stable and accurate representation of sounds despite intervening head movements, the acoustic input should be continuously combined with feedback signals about changes in head orientation. Alternatively, the auditory target coordinates could be updated in advance by using either the preprogrammed gaze-motor command or the sensory target coordinates to which the intervening gaze shift is made ("predictive remapping"). So far, previous experiments cannot dissociate these alternatives. Here, we study whether the auditory system compensates for ongoing saccadic eye and head movements in two dimensions that occur during target presentation. In this case, the system has to deal with dynamic changes of the acoustic cues as well as with rapid changes in relative eye and head orientation that cannot be preprogrammed by the audiomotor system. We performed visual-auditory double-step experiments in two dimensions in which a brief sound burst was presented while subjects made a saccadic eye-head gaze shift toward a previously flashed visual target. Our results show that localization responses under these dynamic conditions remain accurate. Multiple linear regression analysis revealed that the intervening eye and head movements are fully accounted for. Moreover, elevation response components were more accurate for longer-duration sounds (50 msec) than for extremely brief sounds (3 msec), for all localization conditions. Taken together, these results cannot be explained by a predictive remapping scheme. Rather, we conclude that the human auditory system adequately processes dynamically varying acoustic cues that result from self-initiated rapid head movements to construct a stable representation of the target in world coordinates. This signal is subsequently used to program accurate eye and head localization responses.

Acoustic Stimulation↗
PubMed1980

Interaural time differences: implications regarding the neurophysiology of sound localization.

Interaural time differences (ITDs) were measured from 400--7000 Hz on cats in order to provide quantitative data for use in physiological/behavioral studies on sound localization. ITDs derived from clicks and the initial portion of tone bursts showed a pronounced roughness and frequency dependence. This frequency dependence is most evident at higher angles of incidence and indicates that a single ITD will not always represent a single position on the azimuth. Controls demonstrate that most of the roughness in these functions was due to reflections off the surface supporting the animal and that the measured ITDs corresponded to predictions made by steady-state theory. Measurements made with and without the pinnae in position indicate that they have relatively little effect on these ITD functions, particularly for frequencies below 2500 Hz and for small angles of incidence. In spite of acoustic limitations exemplified by the roughness and frequency dependence of these functions, ITDs generated by sound sources situated close to the midline provide reliable localization cues that are much better than those derived from sources well out on the azimuth. Finally, it is noted that another ITD, the group ITD, can be ascribed to an acoustic signal. Calculations based on the measured steady-state ITDs show differences between the group and steady-state ITDs over a given range of frequencies. Differences between the group and steady-state ITD can be significant, and it is argued that: (1) The group ITD can provide a localization cue to the auditory system that is distinct from the steady-state ITD; and (2) it is possible these group ITDs are used by the nervous system to localize sound sources in realistic situations.

Acoustic Stimulation↗
PubMed1986

Effects of auditory cortical lesions on pure-tone sound localization by the albino rat.

Two-choice and three-choice tests were used to evaluate the effects of bilateral auditory cortical lesions on pure-tone sound localization by the albino rat. Both tests required that animals approach a distant sound source to obtain water reinforcement. Stimuli were single noise and tone bursts, 65 ms in duration including 20-ms rise and fall times. Tone frequencies were 2, 4, 8, 16, and 32 kHz adjusted to 40 dB (SPL) above the rat's absolute threshold. Five animals were tested in the two-choice situation following bilateral ablation of auditory cortex. Some reduction in performance was observed relative to normals, but impairments were not severe. Similar results were obtained for 2 brain-damaged animals tested in the three-choice situation. Thus, the ability to localize sounds in space remained intact after complete destruction of auditory cortex, and there was no indication of a frequency-dependent deficit.

Animals↗
PubMed2004

Sound localization in a small passerine bird: discrimination of azimuth as a function of head orientation and sound frequency.

Sound localization is critical to communication when signalers are distributed widely in space and when reverberations that accumulate over distance might otherwise degrade temporal patterns in vocalizations. We readdress the accuracy with which a small passerine bird, the eastern towhee, Pipilo erythrophthalmus L., is able to resolve azimuth in the field. We then report results from two-alternative forced-choice (2AFC) experiments in which three of four subjects were able to discriminate an estimated speaker separation angle of approximately 7 degrees . Subjects oriented laterally when discriminating azimuth in the 2AFC task and each subject preferred a different head orientation. Side biases occurred as a function of head orientation and, as a consequence, we conducted a second 2AFC experiment in which subjects were required to discriminate between two closely spaced lights. Subjects oriented similarly in this visual task, however, side biases did not occur as a function of head orientation. Despite side biases in the auditory task, performance generally declined when subjects were played tones with frequencies near approximately 3 kHz.

Animal Communication↗