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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↗
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↗
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↗
PubMed1992

Hearing in large mammals: sound-localization acuity in cattle (Bos taurus) and goats (Capra hircus).

Sound localization acuity of 3 cattle (Bos taurus) and 2 goats (Capra hircus) was determined for brief complex sounds in a two-choice procedure. Thresholds around the median sagittal plane averaged 30 degrees and 18 degrees, respectively. For comparison, thresholds were obtained in the same test apparatus for humans (0.8 degrees) and a dog (8 degrees). Although the relatively poor acuity of cattle and goats compared with most mammals comes as some surprise, given their large interaural distances and the large binaural locus cues available to them, it is not unexpected when other factors are considered. Like other poor localizers (both domesticated and nondomesticated), cattle and goats are prey species with their best vision directed throughout nearly the entire horizon. In contrast to mammals with very narrow foveal fields, they may not need very accurate locus information from their auditory systems to direct their gaze to a sound source.

Adult↗
PubMed1990

Comparative physiology of sound localization in four species of owls.

Bilateral ear asymmetry is found in some, but not all, species of owls. We investigated the neural basis of sound localization in symmetrical and asymmetrical species, to deduce how ear asymmetry might have evolved from the ancestral condition, by comparing the response properties of neurons in the external nucleus of the inferior colliculus (ICx) of the symmetrical burrowing owl and asymmetrical long-eared owl with previous findings in the symmetrical great horned owl and asymmetrical barn owl. In the ICx of all of these owls, the neurons had spatially restricted receptive fields, and auditory space was topographically mapped. In the symmetrical owls, ICx units were not restricted in elevation, and only azimuth was mapped in ICx. In the barn owl, the space map is two-dimensional, with elevation forming the second dimension. Receptive fields in the long-eared owl were somewhat restricted in elevation, but their tuning was not sharp enough to determine if elevation is mapped. In every species, the primary cue for azimuth was interaural time difference, although ICx units were also tuned for interaural intensity difference (IID). In the barn owl, the IIDs of sounds with frequencies between about 5 and 8 kHz vary systematically with elevation, and the IID selectivity of ICx neurons primarily encodes elevation. In the symmetrical owls, whose ICx neurons do not respond to frequencies above about 5 kHz, IID appears to be a supplementary cue for azimuth. We hypothesize that ear asymmetry can be exploited by owls that have evolved the higher-frequency hearing necessary to generate elevation cues. Thus, the IID selectivity of ICx neurons in symmetrical owls may preadapt them for asymmetry; the neural circuitry that underlies IID selectivity is already present in symmetrical owls, but because IID is not absolutely required to encode azimuth it can come to encode elevation in asymmetrical owls.

Animals↗
PubMed1992

Neural network models of sound localization based on directional filtering by the pinna.

Three-layer neural-network functions were developed to transform spectral representations of pinna-filtered stimuli at the input to a space-mapped representation of sound-source direction at the output. The inputs are modeled after transfer functions of the external ear of the cat; the output is modeled on the spatial sensitivity of superior colliculus neurons. Network solutions are obtained by backpropagation and by a method that enforces uniform task distribution in the hidden layer of the model. Solutions are characterized using bandlimited inputs to study the relative strength of potential sound localization cues in various frequency regions. This analysis suggests that the frequency region containing the first spectral notch (5-18 kHz) provides the best localization cues. Response properties of model neurons were studied using input patterns modeled after auditory nerve response profiles to pure tones at various frequencies and sound levels. The response properties of hidden layer model neurons resemble cochlear nucleus types III and IV and their composites. Neurons in both hidden and output layers show the properties of spectral notch detectors. Although neural networks have limitations as models of real neural systems, the results illustrate how they can provide insight into the computation of complex transformations in the nervous system.

Auditory Perception↗
PubMed1992

Simulated motion enhances neuronal selectivity for a sound localization cue in background noise.

In nature, sound sources move and signals are accompanied by background noise. Noting that motion helps the perception of visual stimuli, we tested whether motion similarly facilitates the detection of acoustic targets, at the neuronal level. Auditory neurons in the central nucleus of the barn owl's inferior colliculus (ICc), due to their selectivity for interaural phase difference (delta phi), are sharply tuned to the azimuth of sound sources and are arrayed to form a topographic map of delta phi. While recording from single ICc neurons, we presented tones that simulated either moving or stationary sound sources with and without background noise. We found that the tuning of cells in the ICc for delta phi was sharper for stimuli that simulated motion than for those that simulated stationary targets. The neurons signaled the presence of a tone obscured by noise better if the tone moved than if the tone remained stationary. The resistance to noise observed with moving stimuli could not be reproduced with the temporal modulation of the stimulus amplitude, suggesting that a change of position over time was required.

Acoustic Stimulation↗
PubMed1992

Pinna-based spectral cues for sound localization in cat.

The directional dependence of the transfer function from free field plane waves to a point near the tympanic membrane (TM) was measured in anesthetized domestic cats. A probe tube microphone was placed approximately 3 mm from the TM from beneath the head in order to keep the pinna intact. Transfer functions were computed as the ratio of the spectrum of a click recorded near the TM to the spectrum of the click in freefield. We analyze the transfer functions in three frequency ranges: low frequencies (less than 5 kHz) where interaural level differences vary smoothly with azimuth; midfrequencies (5-18 kHz) where a prominent spectral notch is observed; and high frequencies (greater than 18 kHz) where the transfer functions vary greatly with source location. Because no two source directions produce the same transfer function, the spectrum of a broadband sound at the TM could serve as a sound localization cue for both elevation and azimuth. In particular, we show that source direction is uniquely determined, for source directions in front of the cat, from the frequencies of the midfrequency spectral notches in the two ears. The validity of the transfer functions as measures of the acoustic input to the auditory system is considered in terms of models of sound propagation in the ear canal.

Acoustic Stimulation↗
PubMed1991

Sound localization in small birds: absolute localization in azimuth.

Nine small birds of 3 species (Melopsittacus undulatus, Serinus canarius, and Poephila guttata) were trained in an operant procedure to fly to sound sources for food reward. The angle between the 2 sound sources was varied on a session-by-session basis, and threshold (i.e., minimum resolvable angle) was taken as the angle that corresponded to a performance level of 75% correct. In all, thresholds were calculated for pure tones of 5 different frequencies, noise bands of 3 different spectral compositions, and species-specific contact or distance calls recorded from each of the 3 species. Thresholds for both simple and complex stimuli were larger than 25 degrees. There were statistically significant species differences for each stimulus set, but these differences were not correlated with species differences in head size. Birds with 1 ear plugged performed as well as binaural birds in this task. Birds deafened in 1 ear, however, performed at chance.

Animals↗
PubMed1978

Some further observations on the effects of unilateral cortical ablation on sound localization in the cat.

It has previously been shown that unilateral ablation of the whole auditory cortex in the cat disrupts the precedence effect, and also interferes with the ability of the normal animal to discriminate in the Y-maze between a single sound on one side and a double sound consisting of a signal on both left and right sides. The present work has confirmed these effects and has shown that both can be obtained with lesions confined to AI and AII. The "one-versus-two" deficit has invariably been seen in all the animals studied, but a proportion of animals do not show the precedence effect deficit. It has been confirmed that the apparent sucess of some animals can be due to the training effect of the one-versus-two paradigm, as was proposed in the earlier paper; however it has also been shown that this cannot be explanation in all cases. It has been demonstrated that cats are able to localize sound behind them with some success; turning around in the start box to reverse right and left space is therefore a possible strategy for overcoming a unilateral deficit. However, even with the head "fixed" in the forward-facing position, one animal was still able to run well above chance. The size of the lesion does not appear to be a correlate of the performance level.

Animals↗
PubMed1975

Neuroanatomical basis of binaural phase-difference analysis for sound localization: a comparative study.

Four varieties of mammals whose medial superior olives range from large to none at all were tested for their ability to localize single, brief tone pips at various frequencies. Although each animal could localize high-frequency tone pips, their ability to localize middle- and low-frequency tone pips corresponded to the size of their medial superior olive (MSO). Since this latter range of frequencies is the one in which binaural phase-difference cues predominate, this anatomical-behavioral correspondence supports the idea that MSO is the chief binaural time-analyzing center for sound localization.

Animals↗
PubMedDate not supplied

Sound localization with phase audiometry.

A 500 Hz pure tone is presented binaurally with earphones. The tone is adjusted to give a midline impression at confortable loudness level. An electronic variable time delay line unit and a generator for randomizing the shift of the phase lag of the signal to the right or to the left ear, respectively, are used. By shortening the time delay the threshold of the recognition of the phase difference is reached. At every test sequence the tone is first presented in the center of the head without delay and then with delay. The patient runs the test by indicating in which ear the tone is heard. The phase difference is thus gradually reduced from 500 musec down to a threshold value of about 48 musec equal on right and left ears in normal-hearing subjects. The results of phase audiometry on subjects with normal hearing and with different types of hearing losses are in good agreement with the results of sound localization tests in free field. Phase audiometry seems to be of especial value in diagnosing retrocochlear lesions.

Acoustic Stimulation↗
PubMed1992

The dominant role of low-frequency interaural time differences in sound localization.

Two experiments are described in which listeners judge the apparent directions of virtual sound sources-headphone-presented sounds that are processed in order to simulate free-field sounds. Previous results suggest that when the cues to sound direction are preserved by the simulation, the apparent directions of virtual sources are nearly the same as the apparent directions of real free-field sources. In the experiments reported here, the interaural phase relations in the processing algorithms are manipulated in order to produce stimuli in which the interaural time difference cues signal one direction and interaural intensity and pinna cues signal another direction. The apparent directions of these conflicting cue stimuli almost always follow the interaural time cue, as long as the wideband stimuli include low frequencies. With low frequencies removed from the stimuli, the dominance of interaural time difference disappears, and apparent direction is determined primarily by interaural intensity difference and pinna cues.

Adult↗
PubMed1992

Azimuthal sound localization in the European starling (Sturnus vulgaris): I. Physical binaural cues.

The physical measurements reported here test whether the European starling (Sturnus vulgaris) evaluates the azimuth direction of a sound source with a peripheral auditory system composed of two acoustically coupled pressure-difference receivers (1) or of two decoupled pressure receivers (2). A directional pattern of sound intensity in the free-field was measured at the entrance of the auditory meatus using a probe microphone, and at the tympanum using laser vibrometry. The maximum differences in the sound-pressure level measured with the microphone between various speaker positions and the frontal speaker position were 2.4 dB at 1 and 2 kHz, 7.3 dB at 4 kHz, 9.2 dB at 6 kHz, and 10.9 dB at 8 kHz. The directional amplitude pattern measured by laser vibrometry did not differ from that measured with the microphone. Neither did the directional pattern of travel times to the ear. Measurements of the amplitude and phase transfer function of the starling's interaural pathway using a closed sound system were in accord with the results of the free-field measurements. In conclusion, although some sound transmission via the interaural canal occurred, the present experiments support the hypothesis 2 above that the starling's peripheral auditory system is best described as consisting of two functionally decoupled pressure receivers.

Acoustic Stimulation↗