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PubMed1990

Effect of bilateral auditory cortex lesions on sound localization in Japanese macaques.

1. The ability of four Japanese macaques (Macaca fuscata) to localize sound was determined after bilateral ablation of auditory cortex. The animals were given two tests: a "midline" test in which they had to discriminate noise bursts presented from a loudspeaker located to the left from identical noise bursts presented from a loudspeaker located to the right of midline, and a "hemifield" test in which both loudspeakers were located in their right hemifield. 2. Both of the tests were administered by the use of two different behavioral tasks: a conditioned-avoidance task in which the animals were trained to make or break contact with a water spout to indicate the location of a sound source, and a two-choice task that required the animals to walk to the source of the sound. 3. The results of both the conditioned-avoidance and the two-choice tasks demonstrated that the animals were able to perform the midline discrimination although their localization acuity was reduced. However, the animals had great difficulty in learning to walk to the source of a sound in spite of the fact that they had received previous sound-localization training in the conditioned-avoidance task. This difficulty suggested that the monkeys no longer associated the sound with a location in space. 4. The results of both the conditioned-avoidance and the two-choice tasks demonstrated that the animals were unable to discriminate the locus of a sound source when both loudspeakers were located in the same hemifield. 5. Bilateral ablation of auditory cortex results in both sensory and perceptual deficits. The presence of sensory deficits is indicated by the decreased acuity in the left-right discrimination and the inability to discriminate between two loudspeakers located in the same hemifield. The deficit in the perception of the locus of sound is indicated by the difficulty in learning to approach the source of a sound, an ability which normal monkeys exhibit without training. 6. There appear to be species' differences in the effect of auditory cortex lesions on sound localization. Although cortical lesions result in a sound-localization deficit in several species of primates and carnivores, they have little or no effect on rats.

Animals↗
PubMed2001

Differential effect of near-threshold stimulus intensities on sound localization performance in azimuth and elevation in normal human subjects.

The ability of humans to localize sounds remains relatively constant across a range of intensities well above detection threshold, and increasing the spectral content of the stimulus results in an improvement in localization ability. For broadband stimuli, intensities near detection threshold result in fewer and weaker binaural cues used in azimuth localization because the stimulus energy at the high- and low-frequency ends of the audible spectrum fall below detection threshold. Thus, the ability to localize broadband sounds in azimuth is predicted to be degraded at audible but near threshold stimulus intensities. The spectral cues for elevation localization (spectral peaks and notches generated by the head-related transfer function) span a narrower frequency range than those for azimuth. As the stimulus intensity decreases, the ability to detect the stimulus frequencies corresponding to the spectral notches will be more strongly affected than the ability to detect frequencies outside the range where these spectral cues are useful. Consequently, decreasing the stimulus intensity should degrade localization in both azimuth and elevation and create a greater deficit in elevation localization due to the narrower band of audible frequencies containing elevation cues compared to azimuth cues. The present study measured the ability of 11 normal human subjects to localize broadband noise stimuli along the midsagittal plane and horizontal meridian at stimulus intensities of 14, 22, and 30 dB above the subject's detection threshold using a go/no-go behavioral paradigm. Localization ability decreased in both azimuth and elevation with decreasing stimulus intensity, and this effect was greater on localization in elevation than on localization in azimuth. The differential effects of stimulus intensity on sound localization in azimuth and elevation found in the present study may provide a valuable tool in investigating the neural correlates of sound location perception.

Adult↗
PubMed2003

[Sound localization cues of binaural hearing].

The ability to localize sound sources in space is of considerable importance to the human safety- and survival-system. Consequently the current scientific interest in improving the safety-standard i. e. in air-traffic control has provided a new momentum for investigating spatial hearing. This review deals with the nature and the relative salience of the localization cues. Localization refers to judgements of the direction and distance of a sound source but here we will deal with direction only. We begin with a short introduction into the so-called Duplex theory which dates back to John William Strutt (later Lord Rayleigh). The idea is that sound localization is based on interaural time differences (ITD) at low frequencies and interaural level differences (ILD) at high frequencies. If the head remains stationary neither a given ITD nor an ILD can sufficiently define the position of a sound source in space. On such a theoretical basis cones of confusion which open outward from each ear can be predicted ambiguously projecting any source on the surface of such a cone onto an interaural axis. Our restricted ability at localizing sound sources in the vertical median plane is another example of possible ambiguity. At the end of the 19th century scientists already realized that occlusion of the pinnae cavities decreases localization competence. As a result of later achievements in physics and signal-theory it became more obvious that the pinnae may provide an additional cue for spatial hearing and that the outer ear together with the head and the upper torso form a sophisticated direction-dependent filter. The action of such a filter is mathematically described by the so-called Anatomical Transfer Function (ATF). The spectral patterning of the sound produced by the pinnae and the head is most effective when the source has spectral energy over a wide range and contains frequencies above 6 kHz, that is it contains wavelengths short enough to interact with the anatomical characteristics of the outer ears. Scientific findings further suggest that spectral patterns like peaks and notches may also be exploited monaurally, albeit an a priori-knowledge at the central-auditive level concerning the corresponding transfer functions and relevant real-world sounds is required. Binaural spectral cues are more likely to play a major role in localization. They are derived from another transfer function, the so-called Interaural Transfer Function (ITF), being the ratio of the ATFs at the two ears. The contributions of all these cues may sometimes not be enough to prevent the listener from opting for the wrong direction. But things can be eased by allowing head-movements: More than 60 years ago science reasoned that small head movements could provide the information necessary to resolve most of the ambiguities. Recent studies have proved that these findings have been accurate all along.

Cues↗
PubMed1998

Kainic acid lesions of the superior olivary complex: effects on sound localization by the albino rat.

The ability of rats to localize sounds in space was determined before and after kainic acid lesions of the superior olivary complex (SOC). Animals were tested with a 45-ms noise burst delivered from loudspeakers on the right or left of midline. Anatomical data showed that the lesions destroyed neurons in SOC while preserving fibers of passage in the trapezoid body and other decussating pathways of the auditory brainstem. Animals with either unilateral or bilateral SOC lesions were impaired in their ability to localize a single noise burst postoperatively. Deficits were also found after unilateral lesions restricted primarily to the lateral superior olive. SOC lesions resulted in an elevation in minimum audible angles for sound localization.

Animals↗
PubMed2000

Sound localization in hemispherectomized subjects: the contribution of crossed and uncrossed cortical afferents.

The aim of the present study was to evaluate how hemispherectomized subjects localize sounds in free field using residual auditory structures under monaural testing conditions. The main objective of using a monaural condition with these subjects, who lack the terminal fields of auditory projections on one side, was to evaluate how the crossed and uncrossed pathways compare, with the aim of resolving this biologically critical function. In this model, crossed and uncrossed inputs refer to auditory stimulation presented to the unobstructed ear on the contralateral and the ipsilateral side of the intact hemisphere, respectively. Three hemispherectomized subjects (Hs) and ten control subjects (Cs) were tested for their accuracy to localize broad band noise bursts (BBNBs) of fixed intensity presented on the horizontal plane. BBNBs were delivered randomly through 16 loudspeakers mounted at 10 degrees intervals on a calibrated perimeter frame located inside an anechoic chamber. Subjects had to report the apparent stimulus location by pointing to its perceived position on the perimeter. Hs were less accurate than Cs in the baseline binaural condition, confirming the finding that with a single hemisphere and/or residual (subcortical) structures they cannot analyze binaural cues to sound localization as efficiently as with two fully functional hemispheres. In the monaural condition, Hs localized poorly when they had to depend on the uncrossed input, but performed as well or even better than the Cs with the crossed input. These findings suggest that monaural spectral cues, which constitute the only residual cue to localization under the monaural testing condition, are treated more efficiently, that is, they lead to better localization performance when relayed to the cortex via crossed pathways than through uncrossed pathways.

Adolescent↗
PubMed1990

An in vitro analysis of sound localization mechanisms in the gerbil lateral superior olive.

One way in which animals localize sounds along the horizon is by detecting the level differences at the 2 ears. Neurons in the lateral superior olive (LSO) encode this cue by integrating the synaptic drive from ipsilateral excitatory and contralateral inhibitory connections. This synaptic integration was analyzed in 400-500-microns brain slices through the gerbil superior olive. Intracellular recordings from LSO neurons were obtained during the application of independent or conjoint electrical stimuli to the excitatory afferent and inhibitory afferent pathways. Stimulation of ascending fibers from the ipsilateral cochlear nucleus reliably evoked EPSPs and action potentials. Stimulation of the medial nucleus of the trapezoid body (MNTB) consistently evoked IPSPs. The evoked postsynaptic potentials differed in that IPSPs were 2 times the duration of EPSPs. An electrophysiological estimate of convergence indicated approximately 10 excitatory and 8 inhibitory afferents per LSO neuron. MNTB stimulation suppressed synaptically evoked action potentials. When stimulus amplitude was increased to the excitatory pathway, it was generally found that a greater MNTB stimulus was necessary to suppress the action potential. A similar commensurate rise in ipsilateral and contralateral acoustic stimulation was also found to be necessary to give the same criterion response. These results confirm that the LSO can integrate evoked action potentials and IPSPs to encode interaural level. Increasing stimulus voltage was found to decrease both action potential and IPSP latency, suggesting that intensity information may be encoded with temporal cues in the nervous system. It was also found that an evoked burst of action potentials could be inhibited in such a way as to yield intermediate discharge rates, dependent on contralateral stimulus level. Taken together, these results suggest that certain properties related to level-difference coding may be available for intracellular analysis using the brain-slice preparation. Several temporal characteristics of the synaptic potentials, including latency and duration, may play a critical role in this simple computation.

Acoustic Stimulation↗
PubMed1994

Sound localization in chinchillas. I: Left/right discriminations.

The ability of chinchillas to localize sound was examined behaviorally using a conditioned avoidance procedure in which the animals were trained to discriminate left from right sound sources. Their minimum audible angle was 15.6 degrees for 100-ms broadband noise making them one of the more accurate rodents, although they are not as accurate as primates and carnivores. Thresholds obtained for filtered noise stimuli demonstrated that chinchillas are equally accurate in localizing either low- or high-frequency noise. Further, they are able to use both interaural phase-difference and interaural intensity-difference cues as demonstrated by their ability to localize both low- and high-frequency pure tones. Finally, analysis of the chinchilla retina supports the hypothesis that the role of auditory localization in directing the eyes to sound sources played a role in the evolution of auditory spatial perception.

Acoustic Stimulation↗
PubMed1999

Sound localization in noise in normal-hearing listeners.

The ability to localize a click train in the frontal-horizontal plane was measured in quiet and in the presence of a white-noise masker. The experiment tested the effects of signal frequency, signal-to-noise ratio (S/N), and masker location. Clicks were low-pass filtered at 11 kHz in the broadband condition, low-pass filtered at 1.6 kHz in the low-pass condition, and bandpass filtered between 1.6 and 11 kHz in the high-pass condition. The masker was presented at either -90, 0, or +90 deg azimuth. Six signal-to-noise ratios were used, ranging from -9 to +18 dB. Results obtained with four normal-hearing listeners show that (1) for all masker locations and filtering conditions, localization accuracy remains unaffected by noise until 0-6 dB S/N and decreases at more adverse signal-to-noise ratios, (2) for all filtering conditions and at low signal-to-noise ratios, the effect of noise is greater when noise is presented at +/- 90 deg azimuth than at 0 deg azimuth, (3) the effect of noise is similar for all filtering conditions when noise is presented at 0 deg azimuth, and (4) when noise is presented at +/- 90 deg azimuth, the effect of noise is similar for the broadband and high-pass conditions, but greater for the low-pass condition. These results suggest that the low- and high-frequency cues used to localize sounds are equally affected when noise is presented at 0 deg azimuth. However, low-frequency cues are less resistant to noise than high-frequency cues when noise is presented at +/- 90 deg azimuth. When both low- and high-frequency cues are available, listeners base their decision on the cues providing the most accurate estimation of the direction of the sound source (high-frequency cues). Parallel measures of click detectability suggest that the poorer localization accuracy observed when noise is at +/- 90 deg azimuth may be caused by a reduction in the detectability of the signal at the ear ipsilateral to the noise.

Adult↗
PubMed1982

Vertical and horizontal sound localization in primates.

Minimum audible angles for localization in the vertical and horizontal planes were psychophysically determined in Old World monkeys (Macaca). In the vertical condition the test stimuli consisted of primate vocalizations and bands of noise. Minimum audible angles ranged from 3 degrees to greater than 20 degrees for signals of various bandwidths. The acuity of vertical localization was dependent upon the high-frequency content of the signal. Two of the three monkeys were unable to vertically localize sounds if the high-frequency limit of the signal was below 2000 Hz. The acuity of horizontal localization was tested for pure tones 500, 2000, and 8000 Hz in frequency, positioned at several referent locations in the right frontal quadrant. The acuity of horizontal localization decreased as the test azimuth was displaced from the midline. Minimum audible angles ranged from approximately 4 degrees to greater than 20 degrees and varied as a function of test azimuth.

Animals↗
PubMed1994

Monaural sound localization: acute versus chronic unilateral impairment.

We tested the ability of human listeners to localize broadband noise bursts in the absence of binaural localization cues. The subject population consisted of five patients, who had normal hearing in one ear and congenital deafness in the other, and seven normal controls, who were tested with both ears open and with one ear plugged. Consistent with previous reports, the introduction of an ear plug unilaterally into control subjects resulted in a prominent lateral displacement in their localization judgements by an average of 30.9 degrees toward the side of the open ear. Vertical localization was less strongly impaired. The five monaural patients showed a considerable range of ability to localize sounds. Two of the patients were essentially indistinguishable from the plugged control subjects in that they showed a prominent displacement of responses toward the side of the hearing ear. The other three subjects localized significantly better than the plugged controls, in that they demonstrated little or no lateral displacement toward the hearing side and that they localized targets on the hearing and on the impaired sides about equally well. The performance of these latter patients demonstrates that monaural cues can provide useful localization information in the horizontal as well as in the vertical dimension.

Acoustic Stimulation↗
PubMed1992

Infants' perception of illusions in sound localization: reaching to sounds in the dark.

Sixteen infants each at 4, 6, and 8 months of age were tested for reaching to sounding toys in the dark under two auditory illusion conditions: the Haas-effect, which creates the illusion of a single lateralized sound based on an interaural intensity difference (the toy was visible and invisible under some test conditions); and the midline illusion, which creates the illusion of a single sound at midline due to an absence of any interaural time or intensity differences (invisible toy condition only). No-sound control trials indicated the level of spontaneous reaching in the dark. Results indicate that by 4 months infants perceive both the Haas-effect and midline illusions. The ability to reach both for invisible and visible sounding objects in the dark was well developed by 4 months of age, although developmental changes in aspects of reaching behavior were observed and, at all ages, object contact was most frequent when visual localization cues accompanied sound localization cues. The incidence of spontaneous reaching in the dark was low and did not vary with age. Theoretical and methodological implications of this research are discussed.

Acoustic Stimulation↗
PubMed1996

Sound localization after kainic acid lesions of the dorsal nucleus of the lateral lemniscus in the albino rat.

The ability of rats to localize sounds in space was determined before and after kainic acid lesions of the dorsal nucleus of the lateral lemniscus (DNLL). The rats were trained to approach a 45-ms noise burst delivered from loudspeakers on the right or left of midline. Lesions were made by local injection of kainic acid into the DNLL. Rats with unilateral lesions of DNLL were impaired in their postoperative ability to localize a single noise burst. Rats with bilateral lesions also had deficits in postoperative performance, but the severity of the impairment was not substantially greater than that expected from a unilateral lesion. The mean pre- and postoperative minimum audible angles were 14.8 degrees and 40.4 degrees for rats with complete unilateral lesions and 13.5 degrees and 36.0 degrees for rats with bilateral lesions.

Acoustic Stimulation↗
PubMed1987

Contribution of auditory cortex to sound localization by the ferret (Mustela putorius).

Ferrets were tested in a semicircular apparatus to determine the effects of auditory cortical lesions on their ability to localize sounds in space. They were trained to initiate trials while facing forward in the apparatus, and sounds were presented from one of two loudspeakers located in the horizontal plane. Minimum audible angles were obtained for three different positions, viz., the left hemifield, with loudspeakers centered around -60 degrees azimuth; the right hemifield, with loudspeakers centered around +60 degrees azimuth; and the midline with loudspeakers centered around 0 degrees azimuth. Animals with large bilateral lesions had severe impairments in localizing a single click in the midline test. Following complete destruction of the auditory cortex performance was only marginally above the level expected by chance even at large angles of speaker separation. Severe impairments were also found in localization of single clicks in both left and right lateral fields. In contrast, bilateral lesions restricted to the primary auditory cortex resulted in minimal impairments in midline localization. The same lesions, however, produced severe impairments in localization of single clicks in both left and right lateral fields. Large unilateral lesions that destroyed auditory cortex in one hemisphere resulted in an inability to localize single clicks in the contralateral hemifield. In contrast, no impairments were found in the midline test or in the ipsilateral hemifield. Unilateral lesions of the primary auditory cortex resulted in severe contralateral field deficits equivalent to those seen following complete unilateral destruction of auditory cortex. No deficits were seen in either the midline or the ipsilateral tests.

Animals↗
PubMed2006

The spatio-temporal brain dynamics of processing and integrating sound localization cues in humans.

Interaural intensity and time differences (IID and ITD) are two binaural auditory cues for localizing sounds in space. This study investigated the spatio-temporal brain mechanisms for processing and integrating IID and ITD cues in humans. Auditory-evoked potentials were recorded, while subjects passively listened to noise bursts lateralized with IID, ITD or both cues simultaneously, as well as a more frequent centrally presented noise. In a separate psychophysical experiment, subjects actively discriminated lateralized from centrally presented stimuli. IID and ITD cues elicited different electric field topographies starting at approximately 75 ms post-stimulus onset, indicative of the engagement of distinct cortical networks. By contrast, no performance differences were observed between IID and ITD cues during the psychophysical experiment. Subjects did, however, respond significantly faster and more accurately when both cues were presented simultaneously. This performance facilitation exceeded predictions from probability summation, suggestive of interactions in neural processing of IID and ITD cues. Supra-additive neural response interactions as well as topographic modulations were indeed observed approximately 200 ms post-stimulus for the comparison of responses to the simultaneous presentation of both cues with the mean of those to separate IID and ITD cues. Source estimations revealed differential processing of IID and ITD cues initially within superior temporal cortices and also at later stages within temporo-parietal and inferior frontal cortices. Differences were principally in terms of hemispheric lateralization. The collective psychophysical and electrophysiological results support the hypothesis that IID and ITD cues are processed by distinct, but interacting, cortical networks that can in turn facilitate auditory localization.

Acoustic Stimulation↗
PubMed2002

Binaural weighting of pinna cues in human sound localization.

Human sound localization relies on binaural difference cues for sound-source azimuth and pinna-related spectral shape cues for sound elevation. Although the interaural timing and level difference cues are weighted to produce a percept of sound azimuth, much less is known about binaural mechanisms underlying elevation perception. This problem is particularly interesting for the frontal hemifield, where binaural inputs are of comparable strength. In this paper, localization experiments are described in which hearing for each ear was either normal, or spectrally disrupted by a mold fitted to the external ear. Head-fixed saccadic eye movements were used as a rapid and accurate indicator of perceived sound direction in azimuth and elevation. In the control condition (both ears free) azimuth and elevation components of saccadic responses were well described by a linear regression line for the entire measured range. For unilateral mold conditions, the azimuth response components did not differ from controls. The influence of the mold on elevation responses was largest on the ipsilateral side, and declined systematically with azimuth towards the side of the free ear. Near the midsagittal plane the elevation responses were clearly affected by the mold, suggesting a systematic binaural interaction in the neural computation of perceived elevation that straddles the midline. A quantitative comparison of responses from the unilateral mold, the bilateral mold and control condition provided evidence that the fusion process can be described by binaural weighted averaging. Two different conceptual schemes are discussed that could underlie the observed responses.

Acoustic Stimulation↗
PubMed1992

Development of sound localization.

Perceptual sound localization is determined from differences in time and loudness of signals arriving separately at bilateral ears, with the time difference being the more important one. The author examined the development of sound lateralization using a self-recording device developed by Sato. It measures the sensitivity of sound lateralization on the basis of time and loudness differences separately. Subjects were 59 children aged 2 to 18 years and 12 adults who were audiologically and neurologically normal. Sound stimulus was 500 Hz continuous band noise. (1) The testable rate was increased in children older than the age of 4, and all children over 7 years of age could be tested. (2) Children over 4 years of age could lateralize sound on the basis of time difference. Sensitivity to time differences improved rapidly, showing a steep curve, in children from the ages of 4 to 6, and then increased slowly until adulthood. (3) Sensitivity to loudness differences increased and standard deviation decreased with age. There is a possibility that children aged 4 can discriminate loudness difference almost as well as adults.

Adolescent↗
PubMed1997

Representation of sound localization cues in the auditory thalamus of the barn owl.

Barn owls can localize a sound source using either the map of auditory space contained in the optic tectum or the auditory forebrain. The auditory thalamus, nucleus ovoidalis (N.Ov), is situated between these two auditory areas, and its inactivation precludes the use of the auditory forebrain for sound localization. We examined the sources of inputs to the N.Ov as well as their patterns of termination within the nucleus. We also examined the response of single neurons within the N.Ov to tonal stimuli and sound localization cues. Afferents to the N.Ov originated with a diffuse population of neurons located bilaterally within the lateral shell, core, and medial shell subdivisions of the central nucleus of the inferior colliculus. Additional afferent input originated from the ipsilateral ventral nucleus of the lateral lemniscus. No afferent input was provided to the N.Ov from the external nucleus of the inferior colliculus or the optic tectum. The N.Ov was tonotopically organized with high frequencies represented dorsally and low frequencies ventrally. Although neurons in the N.Ov responded to localization cues, there was no apparent topographic mapping of these cues within the nucleus, in contrast to the tectal pathway. However, nearly all possible types of binaural response to sound localization cues were represented. These findings suggest that in the thalamo-telencephalic auditory pathway, sound localization is subserved by a nontopographic representation of auditory space.

Acoustic Stimulation↗
PubMed2005

Underwater hearing and sound localization with and without an air interface.

HYPOTHESIS: Underwater hearing acuity and sound localization are improved by the presence of an air interface around the pinnae and inside the external ear canals. BACKGROUND: Hearing threshold and the ability to localize sound sources are reduced underwater. The resonance frequency of the external ear is lowered when the external ear canal is filled with water, and the impedance-matching ability of the middle ear is significantly reduced due to elevation of the ambient pressure, the water-mass load on the tympanic membrane, and the addition of a fluid-air interface during submersion. Sound lateralization on land is largely explained by the mechanisms of interaural intensity differences and interaural temporal or phase differences. During submersion, these differences are largely lost due to the increase in underwater sound velocity and cancellation of the head's acoustic shadow effect because of the similarity between the impedance of the skull and the surrounding water. METHODS: Ten scuba divers wearing a regular opaque face mask or an opaque ProEar 2000 (Safe Dive, Ltd., Hofit, Israel) mask that enables the presence of air at ambient pressure in and around the ear made a dive to a depth of 3 m in the open sea. Four underwater speakers arranged on the horizontal plane at 90-degree intervals and at a distance of 5 m from the diver were used for testing pure-tone hearing thresholds (PTHT), the reception threshold for the recorded sound of a rubber-boat engine, and sound localization. For sound localization, the sound of the rubber boat's engine was randomly delivered by one speaker at a time at 40 dB HL above the recorded sound of a rubber-boat engine, and the diver was asked to point to the sound source. The azimuth was measured by the diver's companion using a navigation board. RESULTS: Underwater PTHT with both masks were significantly higher for frequencies of 250 to 6000 Hz when compared with the thresholds on land (p <0.0001). No differences were found in the PTHT or the reception threshold for the recorded sound of a rubber-boat engine for dry or wet ear conditions. There was no difference in the sound localization error between the regular mask and the ProEar 2000 mask. CONCLUSIONS: The presence of air around the pinna and inside the external ear canal did not improve underwater hearing sensitivity or sound localization. These results support the argument that bone conduction plays the main role in underwater hearing.

Acoustic Impedance Tests↗