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PubMed1992

Directionality of phase locking in auditory nerve fibers of the leopard frog Rana pipiens pipiens.

A dorsal approach to the eighth nerve and free-field stimulation were used to investigate the effect of sound direction and intensity on phase locking in auditory nerve fibers of the leopard frog Rana pipiens pipiens. Tuning curves of 75 auditory neurons were analyzed (Fig. 2). Amphibian papillar neurons, but not basilar papillar neurons, exhibit significant phase locking to short tone bursts at the characteristic frequency (CF), the degree of phase locking (vector strength) decreasing with the neuron's CF (Figs. 3, 4 and 10E). Vector strength increases with sound pressure level to saturate about 20 dB above threshold, while the preferred firing phase is only slightly affected (Figs. 5 and 6). In contrast, sound direction hardly affects vector strength (Figs. 7, 8, 9A and 10A and C), but has a strong influence on the preferred firing phase (Figs. 7, 8, 9B and C, 10B and D): With respect to anterior tone presentation there are phase lags for ipsilateral and phase leads for posterior and contralateral presentation. Phase differences between both ears show a sinusoidal or cardioid/ovoidal directional characteristic; maximum differences are found with antero-lateral tone presentation (Fig. 11). The directionality of phase locking decreases with the neuron's CF (Fig. 10F) and only slightly changes with sound pressure level (Fig. 12). Thus, phase locking of amphibian papilla neurons can potentially provide intensity-independent information for sound localization.

Acoustic Stimulation↗
PubMed2003

Distributed coding of sound locations in the auditory cortex.

Although the auditory cortex plays an important role in sound localization, that role is not well understood. In this paper, we examine the nature of spatial representation within the auditory cortex, focusing on three questions. First, are sound-source locations encoded by individual sharply tuned neurons or by activity distributed across larger neuronal populations? Second, do temporal features of neural responses carry information about sound-source location? Third, are any fields of the auditory cortex specialized for spatial processing? We present a brief review of recent work relevant to these questions along with the results of our investigations of spatial sensitivity in cat auditory cortex. Together, they strongly suggest that space is represented in a distributed manner, that response timing (notably first-spike latency) is a critical information-bearing feature of cortical responses, and that neurons in various cortical fields differ in both their degree of spatial sensitivity and their manner of spatial coding. The posterior auditory field (PAF), in particular, is well suited for the distributed coding of space and encodes sound-source locations partly by modulations of response latency. Studies of neurons recorded simultaneously from PAF and/or A1 reveal that spatial information can be decoded from the relative spike times of pairs of neurons - particularly when responses are compared between the two fields - thus partially compensating for the absence of an absolute reference to stimulus onset.

Acoustic Stimulation↗
PubMed2004

Involvement of the superior temporal cortex and the occipital cortex in spatial hearing: evidence from repetitive transcranial magnetic stimulation.

The processing of auditory spatial information in cortical areas of the human brain outside of the primary auditory cortex remains poorly understood. Here we investigated the role of the superior temporal gyrus (STG) and the occipital cortex (OC) in spatial hearing using repetitive transcranial magnetic stimulation (rTMS). The right STG is known to be of crucial importance for visual spatial awareness, and has been suggested to be involved in auditory spatial perception. We found that rTMS of the right STG induced a systematic error in the perception of interaural time differences (a primary cue for sound localization in the azimuthal plane). This is in accordance with the recent view, based on both neurophysiological data obtained in monkeys and human neuroimaging studies, that information on sound location is processed within a dorsolateral "where" stream including the caudal STG. A similar, but opposite, auditory shift was obtained after rTMS of secondary visual areas of the right OC. Processing of auditory information in the OC has previously been shown to exist only in blind persons. Thus, the latter finding provides the first evidence of an involvement of the visual cortex in spatial hearing in sighted human subjects, and suggests a close interconnection of the neural representation of auditory and visual space. Because rTMS induced systematic shifts in auditory lateralization, but not a general deterioration, we propose that rTMS of STG or OC specifically affected neuronal circuits transforming auditory spatial coordinates in order to maintain alignment with vision.

Acoustic Stimulation↗
PubMed2003

Spatial and non-spatial auditory short-term memory in patients with temporal-lobe lesion.

Primate auditory systems are divided into at least two different pathways. One refers to objects and the other deals with localization. To investigate auditory spatial and non-spatial short-term memory, we tested patients with unilateral medial temporal lobe lesions including the pole in two tasks involving either sound localization discrimination or auditory object discrimination. The results showed that both left and right temporal lobe lesions impaired spatial short-term memory whereas only lesions on the right affected non-spatial short-term memory. By contrast, the same patients were able to perform the tasks when short interstimulus intervals were used suggesting that short-term memory deficits can not be ascribed to difficulties in perception. These findings document, for the first time, in a neurological population, the functional dissociation between spatial and non-spatial auditory short-term memory that seem to depend on separate neural circuits within the medial temporal lobe.

Acoustic Stimulation↗
PubMed2000

Auditory symmetry analysis.

The study of biological symmetry continues to be an important and active area of research, yet in the hearing sciences there are no established quantitative methods for measuring auditory asymmetries and dissimilarities in threshold tuning curves (i.e. audiograms). Using a paired design and adopting methods from the analysis of fluctuating asymmetry, we describe methods for auditory researchers interested in delineating auditory asymmetries and comparing tuning curves, behavioral or neural. We illustrate the methods using audiograms of the prothoracic T-cell interneuron in a nocturnal katydid (Neoconocephalus ensiger). The results show that 87-92 % of T-cells had right-minus-left threshold asymmetries no larger than expected from measurement error alone. Thus, apart from small random fluctuating asymmetries, T-cell pairs in N. ensiger showed no sensory bias and were bilaterally symmetrical from 5 to 100 kHz. The sensitivity of the methods for detecting tuning curve dissimilarities was confirmed in a sound lateralization paradigm by comparing the 'symmetry' (i.e. similarity) of T-cell tuning curves measured at 0 degrees stimulation with tuning curves measured at 90 degrees stimulation for the same T-cell. The results show that T-cell thresholds measured frontally (0 degrees ) were significantly higher than those measured laterally (90 degrees ), particularly for ultrasonic frequencies. Statistically, the directional shift (increase) in auditory thresholds was detected as a directional asymmetry in T-cell tuning, whose origin and functional significance to an insect behaving normally are discussed. The paper discusses practical considerations for detecting auditory asymmetries and tuning curve dissimilarities in general, and closes by questioning the relevance of auditory symmetry for sound localization in both vertebrates and insects.

Animals↗
PubMedDate not supplied

A three-microphone system for real-time directional analysis: toward a device for environmental monitoring in deaf-blind.

Sound localization is a problem for the hearing impaired, deaf, and deaf-blind. A laboratory prototype of a three-microphone digital system for real-time directional analysis of sound sources that is mounted on eyeglasses has been developed. A cross-correlation algorithm is used for determinations of time and phase differences between the three microphone signals. The equipment was evaluated in a sound-treated test room with noise and speech stimuli from 12 different directions. It has an adequate precision (within approximately +/- 10 degrees) for speech and noise bursts in the frequency range 0.5-4 kHz, and for testing with words in a sound field the error percentage is very close to 0 at 0-degrees azimuth. The robustness of the algorithm in terms of signal-to-noise ratio is documented and reaches about +8 dB in different directions for speech and up to 0 dB for noise bursts. It is concluded that the construction of a robust system for directional analysis with sufficient efficiency for further experiments has been achieved.

Algorithms↗
PubMed2002

Tactile "capture" of audition.

Previous research has demonstrated that the localization of auditory or tactile stimuli can be biased by the simultaneous presentation of a visual stimulus from a different spatial position. We investigated whether auditory localization judgments could also be affected by the presentation of spatially displaced tactile stimuli, using a procedure designed to reveal perceptual interactions across modalities. Participants made left-right discrimination responses regarding the perceived location of sounds, which were presented either in isolation or together with tactile stimulation to the fingertips. The results demonstrate that the apparent location of a sound can be biased toward tactile stimulation when it is synchronous, but not when it is asynchronous, with the auditory event. Directing attention to the tactile modality did not increase the bias of sound localization toward synchronous tactile stimulation. These results provide the first demonstration of the tactile capture of audition.

Adult↗
PubMed1995

Binaural tuning of auditory units in the forebrain archistriatal gaze fields of the barn owl: local organization but no space map.

We identified a region in the archistriatum of the barn owl forebrain that contains neurons sensitive to auditory stimuli. Nearly all of these neurons are tuned for binaural localization cues. The archistriatum is known to be the primary source of motor-related output from the avian forebrain and, in barn owls, contributes to the control of gaze, much like the frontal eye fields in monkeys. The auditory region is located in the medial portion of the archistriatum, at the level of the anterior commissure, and is within the region of the archistriatum from which head saccades can be elicited by electrical microstimulation (see preceding companion article, Knudsen et al., 1995). Free-field measurements revealed that auditory sites have large, spatial receptive fields. However, within these large receptive fields, responses are tuned sharply for sound source location. Dichotic measurements showed that auditory sites are tuned broadly for frequency and that the majority are tuned to particular values of interaural time differences and interaural level differences, the principal cues used by barn owls for sound localization. The tuning of sites to these binaural cues is essentially independent of sound level. The auditory properties of units in the medial archistriatum are similar to those of units in the optic tectum, a structure that also contributes to gaze control. Unlike the optic tectum, however, the auditory region of the archistriatum does not contain a single, continuous auditory map of space. Instead, it is organized into dorsoventral clusters of sites with similar binaural (spatial) tuning. The different representations of auditory space in closely related structures in the forebrain (archistriatum) and midbrain (optic tectum) probably reflect the fact that the forebrain contributes to a wide variety of sensorimotor tasks more complicated than gaze control.

Acoustic Stimulation↗
PubMed2000

Auditory cortical responses to the interactive effects of interaural intensity disparities and frequency.

Under natural conditions, stimuli reaching the two ears contain multiple acoustic components. Rarely does a stimulus containing only one component (e.g. pure tone burst) exist outside the realm of the laboratory. For example, in sound localization the simultaneous presence of multiple cues (spectral content, level, phase, etc.) serves to increase the number of available cues and provide the listener with more information, thereby helping to reduce errors in locating the sound source. The present study was designed to explore the relationship between two acoustic parameters: stimulus frequency and interaural intensity disparities (IIDs). By varying both stimulus frequency and IIDs for each cell, we hoped to gain insight into how multiple cues are processed. To this end, we examined the responses of neurons in cat primary auditory cortex (AI) to determine if their sensitivity to IIDs changed as a function of stimulus frequency. IIDs ranging from +30 to -30 dB were presented at different frequencies (frequency was always the same in the two ears). We found that approximately half of the units examined exhibited responses to IIDs that varied as a function of stimulus frequency (i.e. displayed some form of IID x Freq dependency). The remaining units displayed IID responses that were not clearly related to stimulus frequency.

Acoustic Stimulation↗
PubMed2006

Task-modulated "what" and "where" pathways in human auditory cortex.

Human neuroimaging studies suggest that localization and identification of relevant auditory objects are accomplished via parallel parietal-to-lateral-prefrontal "where" and anterior-temporal-to-inferior-frontal "what" pathways, respectively. Using combined hemodynamic (functional MRI) and electromagnetic (magnetoencephalography) measurements, we investigated whether such dual pathways exist already in the human nonprimary auditory cortex, as suggested by animal models, and whether selective attention facilitates sound localization and identification by modulating these pathways in a feature-specific fashion. We found a double dissociation in response adaptation to sound pairs with phonetic vs. spatial sound changes, demonstrating that the human nonprimary auditory cortex indeed processes speech-sound identity and location in parallel anterior "what" (in anterolateral Heschl's gyrus, anterior superior temporal gyrus, and posterior planum polare) and posterior "where" (in planum temporale and posterior superior temporal gyrus) pathways as early as approximately 70-150 ms from stimulus onset. Our data further show that the "where" pathway is activated approximately 30 ms earlier than the "what" pathway, possibly enabling the brain to use top-down spatial information in auditory object perception. Notably, selectively attending to phonetic content modulated response adaptation in the "what" pathway, whereas attending to sound location produced analogous effects in the "where" pathway. This finding suggests that selective-attention effects are feature-specific in the human nonprimary auditory cortex and that they arise from enhanced tuning of receptive fields of task-relevant neuronal populations.

Adaptation, Physiological↗
PubMed2003

Adaptive plasticity in the auditory thalamus of juvenile barn owls.

Little is known about the capacity of the thalamus for experience-dependent plasticity. Here, we demonstrate adaptive changes in the tuning of auditory thalamic neurons to a major category of sound localization cue, interaural time differences (ITDs), in juvenile barn owls that experience chronic abnormal hearing. Abnormal hearing was caused by a passive acoustic filtering device implanted in one ear that altered the timing and level of sound differently at different frequencies. Experience with this device resulted in adaptive, frequency-dependent shifts in the tuning of thalamic neurons to ITD that mimicked the acoustic effects of the device. Abnormal hearing did not alter ITD tuning in the central nucleus of the inferior colliculus, the primary source of input to the auditory thalamus. Therefore, the thalamus is the earliest stage in the forebrain pathway in which this plasticity is expressed. A visual manipulation, chronic prismatic displacement of the visual field, which causes adaptive changes in ITD tuning at higher levels in the forebrain, had no effect on thalamic ITD tuning. The results demonstrate that, during the juvenile period, auditory experience shapes neuronal response properties in the thalamus in a frequency-specific manner and suggest that this thalamic plasticity is driven by self-organizational forces and not by visual instruction.

Acoustic Stimulation↗
PubMed1990

[Directional hearing in relation to individual circadian biorhythm].

Acuity angle of the directional hearing was investigated in connection with the individual circadian rhythm. Two groups of 15 persons represented the morning and evening form of the circadian rhythm. Body temperature fixed the rhythm character. The evaluations of the angle acuity of the directional hearing were performed in the highest and the lowest point of body temperature as well as in the neutral point, which was determined in the morning group in the middle between the two extremes. The possibility of the sound localization in individual and linked with the body temperature circadian rhythm.

Acoustic Stimulation↗
PubMed1986

An acoustic approach to diver navigation.

Three experiments were carried out to assess the capability of divers to localize acoustic signals underwater and to navigate by them. In the first experiment, divers attempted to determine the correct distance to several underwater transducers projecting acoustic stimuli on a horizontal plane. The second experiment consisted of two related studies where the diver/subjects attempted to discover which of many possible stimuli would produce the most robust perception of underwater "sound movement," or the Underwater Auditory Phi Phenomenon (UAPP). A third experiment consisted of navigational swims by divers; the acoustic stimuli utilized were based on those identified in prior experiments as the most preferred. The results demonstrated that divers are able to discriminate among signals emanating from acoustic sources at various distances underwater and to do so at levels well above chance. Second, divers judged 500-Hz square waves to be the signal which best facilitated an acceptable UAPP; thermal noise and 1-kHz square waves followed in that order. However, these differences were only slight and, in practice, divers maintained that the noise signal was the most useful. Third, it was found that divers apparently can effectively navigate by means of auditory signals alone--at least within certain limits. Finally, a significant decrease in the discrimination abilities of divers for frequencies above 6 kHz suggests that intensity cues may not be as robust as time-of-arrival information with respect to underwater sound localization.

Acoustic Stimulation↗
PubMed2004

A neuronal correlate of the precedence effect is associated with spatial selectivity in the barn owl's auditory midbrain.

Sound localization in echoic conditions depends on a precedence effect (PE), in which the first arriving sound dominates the perceived location of later reflections. Previous studies have demonstrated neurophysiological correlates of the PE in several species, but the underlying mechanisms remain unknown. The present study documents responses of space-specific neurons in the barn owl's inferior colliculus (IC) to stimuli simulating direct sounds and reflections that overlap in time at the listener's ears. Responses to 100-ms noises with lead-lag delays from 1 to 100 ms were recorded from neurons in the space-mapped subdivisions of IC in anesthetized owls (N2O/isofluorane). Responses to a target located at a unit's best location were usually suppressed by a masker located outside the excitatory portion of the spatial receptive field. The least spatially selective units exhibited temporally symmetric effects, in that the amount of suppression was the same whether the masker led or lagged. Such effects mirror the alteration of localization cues caused by acoustic superposition of leading and lagging sounds. In more spatially selective units, the suppression was often temporally asymmetric, being more pronounced when the masker led. The masker often evoked small changes in spatial tuning that were not related to the magnitude of suppressive effects. The association of temporally asymmetric suppression with spatial selectivity suggests that this property emerges within IC, and not at earlier stages of auditory processing. Asymmetric suppression reduces the ability of highly spatially selective neurons to encode the location of lagging sounds, providing a possible basis for the PE.

Acoustic Stimulation↗
PubMed2006

Auditory evoked magnetic fields in relation to interaural time delay and interaural correlation.

The detection of interaural time differences (ITD) for sound localization depends on the similarity between the left and right ear signals, namely interaural correlation (IAC). Human localization performance deteriorates with decreasing IACs. In order to examine activity related to localization performance in the human cortex, auditory evoked magnetic fields to the ITD of bandpass noises with different IACs were analyzed. When the IAC was 0.95, the N1m amplitudes, i.e., the estimated equivalent current dipole moments, increased with increasing ITD. However the effect of ITD on the N1m amplitudes was not significant when the IAC was 0.5. When the ITD was 0.7 ms, the N1m amplitudes decreased with decreasing IACs. There were no systematic changes in the source location of N1m in the auditory cortex related to changes in ITD or IAC. The results suggest that localization performance is reflected in N1m amplitudes.

Acoustic Stimulation↗
PubMed1990

Directional dependence of interaural envelope delays.

Interaural envelope delays were measured in six human subjects as a function of the location of a movable sound source, bandpassed between 3 and 16 kHz. A total of 324 source locations were tested in horizontal and vertical increments of 10 degrees. A method is described for estimating the complex directional transfer function of the external ear, independent of the position of the recording microphone in the ear canal. To compute interaural envelope delays, directional transfer functions from the left and right ears were convolved with a critical-band filter, the envelopes were computed, and the envelopes were cross correlated. Interaural envelope delays, as well as interaural group delays, varied somewhat with the center frequency of the critical-band filter and with the vertical location of the sound source. Nevertheless, to a first approximation, envelope delays measured in the ear canals increased monotonically with increasing angle of incidence relative to the median plane, as they would for two microphones on the surface of a rigid sphere. The results are discussed in relation to the possible contribution of interaural envelope delays to sound localization behavior.

Attention↗
PubMed1992

Neural derivation of sound source location: resolution of spatial ambiguities in binaural cues.

Cues for sound localization are inherently spatially ambiguous. Nevertheless, most neurons in the barn owl's optic tectum (superior colliculus) have receptive fields for broadband noise stimuli that are restricted to a single region of space. This study characterizes the spatial ambiguities associated with two important sets of localization cues, interaural level differences (ILDs) and interaural phase differences (IPDs), and describes how information is integrated within and across frequencies to resolve these ambiguities. The auditory receptive fields of neurons in the optic tectum were measured with free-field sounds presented from a movable loudspeaker. In contrast to the single regions typical for broadband receptive fields, receptive fields for tonal stimuli usually included additional discrete regions of space (accessory fields). Based on acoustic measurements of ILD and IPD cues made in the external ear canals, it was shown that accessory fields corresponded to locations from which sound sources produced ILD and IPD values that were approximately the same as those arising from the broadband receptive field. In addition, accessory fields had inhibitory surrounds, corresponding to locations from which sound sources produced substantially different combinations of ILD and IPD values. Where an accessory field for one frequency overlapped with the inhibitory surround of a second frequency, an excitatory response to the first frequency could be reduced or eliminated by addition of the second frequency. Because tonal receptive fields for different frequencies always overlapped in the region of the broadband receptive field but tended not to overlap elsewhere, this integration of excitation and inhibition can account for the restriction of broadband receptive fields to a single region of space.

Animals↗
PubMed1994

Adaptive plasticity of the auditory space map in the optic tectum of adult and baby barn owls in response to external ear modification.

1. This study demonstrates the influence of experience on the establishment and maintenance of the auditory map of space in the optic tectum of the barn owl. Auditory experience was altered either by preventing the structures of the external ears (the facial ruff and preaural flaps) from appearing in baby barn owls (baby ruff-cut owls) or by removing these structures in adults (adult ruff-cut owls). These structures shape the binaural cues used for localizing sounds in both the horizontal and vertical dimensions. 2. The acoustic effects of removing the external ear structures were measured using probe tube microphones placed in the ear canals. In both baby and adult ruff-cut owls, the spatial pattern of binaural localization cues was dramatically different from normal: interaural level difference (ILD) changed with azimuth instead of with elevation, the rate of change of ILD across space was decreased relative to normal, and the rate of change of interaural time difference (ITD) across frontal space was increased relative to normal. 3. The neurophysiological representations of ITD and ILD in the optic tectum were measured before and > or = 3 mo after ruff removal in adults and beginning at 4.5 months of age in baby ruff-cut owls. Multiunit tuning to ITD and to ILD was measured using dichotic stimulation in ketamine-anesthetized owls. The tectal maps of ITD and ILD were reconstructed using visual receptive field location as a marker for recording site location in the optic tectum. 4. Adjustment of the tectal map of ITD to the altered spatial pattern of acoustic ITD was essentially complete in adults as well as in baby ruff-cut owls. This adjustment changed the magnification of ITD across the tectum, with resultant changes in ITD tuning at individual tectal sites of up to approximately 25 microseconds (approximately 5% of the physiological range) relative to normal values. 5. Adaptation of the tectal ILD map to the ruff-cut spatial pattern of acoustic ILD was substantial but clearly incomplete in both adult and baby ruff-cut owls. Although changes of up to approximately 15 dB (approximately 47% of the physiological range) relative to normal tuning were observed at certain tectal sites, the topography of the ILD map was always intermediate between normal and that predicted by the ruff-cut spatial pattern of acoustic ILD.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗