Interaural Time Disparities Research Articles

Neural representation of interaural correlation in human auditory brainstem: Comparisons between temporal-fine structure and envelope

Central processing of interaural correlation (IAC), which depends on the precise representation of acoustic signals from the two ears, is essential for both localization and recognition of auditory objects. A complex soundwave is initially filtered by the peripheral auditory system into multiple narrowband waves, which are further decomposed into two functionally distinctive components: the quickly-varying temporal-fine structure (TFS) and the slowly-varying envelope. In rats, a narrowband noise can evoke auditory-midbrain frequency-following responses (FFRs) that contain both the TFS component (FFRTFS) and the envelope component (FFREnv), which represent the TFS and envelope of the narrowband noise, respectively. These two components are different in sensitivity to the interaural time disparity. In human listeners, the present study investigated whether the FFRTFS and FFREnv components of brainstem FFRs to a narrowband noise are different in sensitivity to IAC and whether there are potential brainstem mechanisms underlying the integration of the two components. The results showed that although both the amplitude of FFRTFS and that of FFREnv were significantly affected by shifts of IAC between 1 and 0, the stimulus-to-response correlation for FFRTFS, but not that for FFREnv, was sensitive to the IAC shifts. Moreover, in addition to the correlation between the binaurally evoked FFRTFS and FFREnv, the correlation between the IAC-shift-induced change of FFRTFS and that of FFREnv was significant. Thus, the TFS information is more precisely represented in the human auditory brainstem than the envelope information, and the correlation between FFRTFS and FFREnv for the same narrowband noise suggest a brainstem binding mechanism underlying the perceptual integration of the TFS and envelope signals.

A model of the neuronal processing of interaural time disparities

The auditory system of humans and other mammals is able to use interaural time and intensity differences as well as spectral cues to localize sound sources. One important cue for localizing low-frequency sound sources in the horizontal plane are interaural time differences (ITDs), which are first analyzed in the medial superior olive (MSO) in the brainstem. Results from electrophysiological and psychoacoustic studies suggest ITD encoding in the relative activities of neuronal populations in the two brain hemispheres. This contribution first explores the neuronal encoding of fine-structure ITDs using a physiologically motivated spiking neuronal-network model of the mammalian MSO. Results from this model confirm robust ITD encoding in the relative activity of the MSOs in both hemispheres. Based on the neuronal-network simulations, a simple probabilistic model of subsequent ITD processing is proposed. This simplified model connects the neuronal responses of the two hemispheres with different hearing sensation properties, such as lateral position, spatial extent, or number of spatially separable sensations. First predictions from the model are discussed with regard to the results of a series of psychoacoustic experiments on the lateralization of pure-tone impulses presented via headphones.

Neural representations of concurrent sounds with overlapping spectra in rat inferior colliculus: Comparisons between temporal-fine structure and envelope

Perceptual segregation of multiple sounds, which overlap in both time and spectra, into individual auditory streams is critical for hearing in natural environments. Some cues such as interaural time disparities (ITDs) play an important role in the segregation, especially when sounds are separated in space. In this study, we investigated the neural representation of two uncorrelated narrowband noises that shared the identical spectrum in the rat inferior colliculus (IC) using frequency-following-response (FFR) recordings, when the ITD for each noise stimulus was manipulated. The results of this study showed that recorded FFRs exhibited two distinctive components: the fast-varying temporal fine structure (TFS) component (FFRTFS) and the slow-varying envelope component (FFRENV). When a single narrowband noise was presented alone, the FFRTFS, but not the FFRENV, was sensitive to ITDs. When two narrowband noises were presented simultaneously, the FFRTFS took advantage of the ITD disparity that was associated with perceived spatial separation between the two concurrent sounds, and displayed a better linear synchronization to the sound with an ipsilateral-leading ITD. However, no effects of ITDs were found on the FFRENV. These results suggest that the FFRTFS and FFRENV represent two distinct types of signal processing in the auditory brainstem and contribute differentially to sound segregation based on spatial cues: the FFRTFS is more critical to spatial release from masking.

Physiology and anatomy of neurons in the medial superior olive of the mouse

In mammals with good low-frequency hearing, the medial superior olive (MSO) computes sound location by comparing differences in the arrival time of a sound at each ear, called interaural time disparities (ITDs). Low-frequency sounds are not reflected by the head, and therefore level differences and spectral cues are minimal or absent, leaving ITDs as the only cue for sound localization. Although mammals with high-frequency hearing and small heads (e.g., bats, mice) barely experience ITDs, the MSO is still present in these animals. Yet, aside from studies in specialized bats, in which the MSO appears to serve functions other than ITD processing, it has not been studied in small mammals that do not hear low frequencies. Here we describe neurons in the mouse brain stem that share prominent anatomical, morphological, and physiological properties with the MSO in species known to use ITDs for sound localization. However, these neurons also deviate in some important aspects from the typical MSO, including a less refined arrangement of cell bodies, dendrites, and synaptic inputs. In vitro, the vast majority of neurons exhibited a single, onset action potential in response to suprathreshold depolarization. This spiking pattern is typical of MSO neurons in other species and is generated from a complement of Kv1, Kv3, and IH currents. In vivo, mouse MSO neurons show bilateral excitatory and inhibitory tuning as well as an improvement in temporal acuity of spiking during bilateral acoustic stimulation. The combination of classical MSO features like those observed in gerbils with more unique features similar to those observed in bats and opossums make the mouse MSO an interesting model for exploiting genetic tools to test hypotheses about the molecular mechanisms and evolution of ITD processing.

Glycinergic transmission modulates GABAergic inhibition in the avian auditory pathway

For all neurons, a proper balance of synaptic excitation and inhibition is crucial to effect computational precision. Achievement of this balance is remarkable when one considers factors that modulate synaptic strength operate on multiple overlapping time scales and affect both pre- and postsynaptic elements. Recent studies have shown that inhibitory transmitters, glycine and GABA, are co-released in auditory nuclei involved in the computation of interaural time disparities (ITDs), a cue used to process sound source location. The co-release expressed at these synapses is heavily activity dependent, and generally occurs when input rates are high. This circuitry, in both birds and mammals, relies on inhibitory input to maintain the temporal precision necessary for ITD encoding. Studies of co-release in other brain regions suggest that GABA and glycine receptors (GlyRs) interact via cross-suppressive modulation of receptor conductance. We performed in vitro whole-cell recordings in several nuclei of the chicken brainstem auditory circuit to assess whether this cross-suppressive phenomenon was evident in the avian brainstem. We evaluated the effect of pressure-puff applied glycine on synaptically evoked inhibitory currents in nucleus magnocellularis (NM) and the superior olivary nucleus (SON). Glycine pre-application reduced the amplitude of inhibitory postsynaptic currents (IPSCs) evoked during a 100 Hz train stimulus in both nuclei. This apparent glycinergic modulation was blocked in the presence of strychnine. Further experiments showed that this modulation did not depend on postsynaptic biochemical interactions such as phosphatase activity, or direct interactions between GABA and GlyR proteins. Rather, voltage clamp experiments in which we manipulated Cl− flux during agonist application suggest that activation of one receptor will modulate the conductance of the other via local changes in Cl− ion concentration within microdomains of the postsynaptic membrane.

Slowly emerging glycinergic transmission enhances inhibition in the sound localization pathway of the avian auditory system

Localization of low-frequency acoustic stimuli is processed in dedicated neural pathways where coincidence-detecting neurons compare the arrival time of sound stimuli at the two ears, or interaural time disparity (ITD). ITDs occur in the submillisecond range, and vertebrates have evolved specialized excitatory and inhibitory circuitry to compute these differences. Glycinergic inhibition is a computationally significant and prominent component of the mammalian ITD pathway. However, evidence for glycinergic transmission is limited in birds, where GABAergic inhibition has been thought to be the dominant or exclusive inhibitory transmitter. Indeed, previous work showed that GABA antagonists completely eliminate inhibition in avian nuclei specialized for processing temporal features of sound, nucleus magnocellularis (NM) and nucleus laminaris (NL). However, more recent work shows that glycine is coexpressed with GABA in synaptic terminals apposed to neurons in both nuclei (Coleman WL, Fischl MJ, Weimann SR, Burger RM. J Neurophysiol 105: 2405-2420, 2011; Kuo SP, Bradley LA, Trussell LO. J Neurosci 29: 9625-9634, 2009). Here we show complementary evidence of functional glycine receptor (GlyR) expression in NM and NL. Additionally, we show that glycinergic input can be evoked under particular stimulus conditions. Stimulation at high but physiologically relevant rates evokes a slowly emerging glycinergic response in NM and NL that builds over the course of the stimulus. Glycinergic response magnitude was stimulus rate dependent, representing 18% and 7% of the total inhibitory current in NM and NL, respectively, at the end of the 50-pulse, 200-Hz stimulus. Finally, we show that the glycinergic component is functionally relevant, as its elimination reduced inhibition of discharges evoked by current injection into NM neurons.

Inhibitory synaptic release properties are topographically distributed in auditory circuitry

Inhibitory synaptic release properties are topographically distributed in auditory circuitry

Modulation of synaptic input by GABAB receptors improves coincidence detection for computation of sound location

Interaural time disparities (ITDs) are the primary cues for localisation of low-frequency sound stimuli. ITDs are computed by coincidence-detecting neurones in the medial superior olive (MSO) in mammals. Several previous studies suggest that control of synaptic gain is essential for maintaining ITD selectivity as stimulus intensity increases. Using acute brain slices from postnatal day 7 to 24 (P7–P24) Mongolian gerbils, we confirm that activation of GABAB receptors reduces the amplitude of excitatory and inhibitory synaptic currents to the MSO and, moreover, show that the decay kinetics of IPSCs are slowed in mature animals. During repetitive stimuli, activation of GABAB receptors reduced the amount of depression observed, while PSC suppression and the slowed kinetics were maintained. Additionally, we used physiological and modelling approaches to test the potential impact of GABAB activation on ITD encoding in MSO neurones. Current clamp recordings from MSO neurones were made while pharmacologically isolated excitatory inputs were bilaterally stimulated using pulse trains that simulate ITDs in vitro. MSO neurones showed strong selectivity for bilateral delays. Application of both GABAB agonists and antagonists demonstrate that GABAB modulation of synaptic input can sharpen ITD selectivity. We confirmed and extended these results in a computational model that allowed for independent manipulation of each GABAB-dependent effect. Modelling suggests that modulation of both amplitude and kinetics of synaptic inputs by GABAB receptors can improve precision of ITD computation. Our studies suggest that in vivo modulation of synaptic input by GABAB receptors may act to preserve ITD selectivity across various stimulus conditions.

Synaptic plasticity in the medial superior olive of hearing, deaf, and cochlear‐implanted cats

The medial superior olive (MSO) is a key auditory brainstem structure that receives binaural inputs and is implicated in processing interaural time disparities used for sound localization. The deaf white cat, a proven model of congenital deafness, was used to examine how deafness and cochlear implantation affected the synaptic organization at this binaural center in the ascending auditory pathway. The patterns of axosomatic and axodendritic organization were determined for principal neurons from the MSO of hearing, deaf, and deaf cats with cochlear implants. The nature of the synapses was evaluated through electron microscopy, ultrastructure analysis of the synaptic vesicles, and immunohistochemistry. The results show that the proportion of inhibitory axosomatic terminals was significantly smaller in deaf animals when compared with hearing animals. However, after a period of electrical stimulation via cochlear implants the proportion of inhibitory inputs resembled that of hearing animals. Additionally, the excitatory axodendritic boutons of hearing cats were found to be significantly larger than those of deaf cats. Boutons of stimulated cats were significantly larger than the boutons in deaf cats, although not as large as in the hearing cats, indicating a partial recovery of excitatory inputs to MSO dendrites after stimulation. These results exemplify dynamic plasticity in the auditory brainstem and reveal that electrical stimulation through cochlear implants has a restorative effect on synaptic organization in the MSO.

Spatial release from speech-on-speech masking in the median sagittal plane

Several studies have described a release from speech-on-speech masking associated with separation of target and masker sources in the median sagittal plane. Some have excluded the possibility that small differences between target and masker interaural time disparities can fully account for this release. This study explored the mechanisms underlying the spatial release from speech-on-speech masking that can be obtained in the absence of such differences. In one condition, interaural time disparities were removed from the nominal median-sagittal-plane, head-related impulse responses used to generate the virtual auditory space within which competing sentences were presented. In other conditions, interaural level and spectral disparities also were manipulated by presenting competing sentences monaurally or diotically after convolution with one ear's head-related impulse responses. It was found that substantial spatial release from masking can be obtained in the absence of any interaural disparities and that such disparities probably make a relatively minor contribution to spatial release from speech-on-speech masking in the median sagittal plane. It is argued that this release from masking is driven primarily by a reduction in informational masking that occurs when monaural information at one, or both, of the listener's ears facilitates differentiation of competing sentences that emanate from spatially separated sources.

Distinguishing feature misperception from illusory conjunctions in spatially distributed musical tones

Recent questions have been raised in the visual search literature concerning whether illusory conjunctions of correctly registered features occur (indicating a feature integration process) or are an artifact of feature misperception. The current investigation raised and addressed similar questions for findings of auditory illusory conjunctions using simultaneous, spatially distributed musical tones. Two experiments were conducted where musically trained listeners identified pairs of tones that reflected possible combinations of two instrument timbres with two pitches. Experiments differed in the spatial separation between simultaneous tones to potentially evaluate the effects of distance on feature perception/integration. In Experiment 1 tones were presented to opposing ears. In Experiment 2 tones were only slightly lateralized by a manipulation of interaural time disparities. Conjunction responses, reflecting the incorrect combination of features, frequently occurred, and were more common for slightly lateralized tones. To evaluate the perceptual event(s) responsible for conjunction responses, data were submitted to multinomial models that differed with respect to whether or not they allowed for illusory conjunctions, the misperception of features, or both errors. Across experiments, data fitting by models was improved by feature misperception, but was not further improved by illusory conjunctions. Implications for models of search performance and feature binding are discussed.

Evidence for efferent effects on early components of the human auditory brain-stem evoked potentials

Objectives and methods: Auditory brain-stem evoked potentials (ABEPs) were recorded from 10 normal hearing subjects in response to rarefaction clicks, presented at a rate of 11/s. Stimuli were binaurally symmetrical and isochronic at 75 dB peSPL or with interaural time disparities (ITDs) of ±0.4 ms, or intensity disparities (IIDs) of ±10 dB. Potentials were recorded from vertex-neck, as well as from 3 orthonormally positioned differential derivations. The amplified potentials were averaged over 8000 repetitions using a dwell time of 20 μs/address/channel. The effects of contralateral stimulation on neural responses of the peripheral auditory system were obtained by subtracting the binaural response from the algebraic sum of responses to left and right monaural stimuli. From the 3 orthonormal derivations, 3-channel Lissajous' trajectories (3-CLTs) to the various stimulus conditions and difference waveforms were derived. Results: The results corroborated earlier studies on binaural interaction components (BICs), which include 3 major components corresponding in latency to the vertex-mastoid peaks IV–VI of ABEP. In addition, the binaural difference waveforms included 3 earlier, low-amplitude components. Latency correspondence and comparison of difference waveform and ABEP 3-CLTs indicated that the first and third early difference waveform components corresponded to the negative peaks following I and III, respectively, of the vertex-neck ABEP to binaural clicks. Conclusions: These results indicate that early ABEP peaks, generated peripheral to binaural convergence, may be affected by contralateral stimulation. These contralateral effects were in a pattern compatible with suppression. most probably by efferents of the olivo-cochlear bundle.

Effects of localized pontine lesions on auditory brain-stem evoked potentials and binaural processing in humans

Objectives and mehtods: Four sets of measurements were obtained from 11 patients (44–80 years old) with small, localized pontine lesions due to vascular disease: (1) Monaural auditory brain-stem evoked potentials (ABEPs; peaks I to VI); (2) Binaural ABEPs processed for their binaural interaction components (BICs) in the latency range of peaks IV to VI; (3) magnetic resonance imaging (MRI) of the brain-stem; and (4) psychoacoustics of interaural time disparity measures of binaural localization. ABEPs and BICs were analyzed for peak latencies and interpeak latency differences. Three-channel Lissajous' trajectories (3-CLTs) were derived for ABEPs and BICs and the latencies and orientations of the equivalent dipoles of ABEP and BICs were inferred from them. Results: Intercomponent latency measures of monaurally evoked ABEPs were abnormal in only 3 of the 11 patients. Consistent correlations between sites of lesion and neurophysiological abnormality were obtained in 9 of the 11 patients using 3-CLT measures of BICs. Six of the 11 patients had absence of one or more BIC components. Seven of the 11 had BICs orientation abnormality and 3 had latency abnormalities. Trapezoid body (TB) lesions (6 patients) were associated with an absent (two patients with ventral-caudal lesions) or abnormal (one patient with ventral-rostral lesions) dipole orientation of the first component (at the time of ABEPs IV), and sparing of this component with midline ventral TB lesions (two patients). A deviant orientation of the second BICs component (at the time of ABEPs V) was observed with ventral TB lesions. Psychoacoustic lateralization in these patients was biased toward the center. Rostral lateral lemniscus (LL) lesions (3 patients) were associated with absent (one patient) or abnormal (two patients) orientation of the third BICs component (at the time of ABEPs VI); and a side-biased lateralization with behavioral testing. Conclusions: These results indicate that: (1) the BICs component occurring at the time of ABEPs peak IV is dependent on ventral-caudal TB integrity; (2) the ventral TB contributes to the BICs component at the time of ABEPs peak V; and (3) the rostral LL is a contributing generator of the BICs component occurring at the time of ABEP peak VI.

Features of ipsilaterally evoked inhibition in the dorsal nucleus of the lateral lemniscus

The dorsal nucleus of the lateral lemniscus (DNLL) is a binaural nucleus whose neurons are excited by stimulation of the contralateral ear and inhibited by stimulation of the ipsilateral ear. Here we report on several features of the ipsilaterally evoked inhibition in 95 DNLL neurons of the mustache bat. These features include its dependence on intensity, its tuning and the types of stimuli that are capable of evoking it. Inhibition was studied by evoking discharges with the iontophoretic application of glutamate, and then evaluating the strength and duration of the inhibition of the glutamate evoked background activity produced by stimulation of the ipsilateral ear. Excitatory responses were evoked by stimulation of the contralateral ear with best frequency (BF) tone bursts. Glutamate evoked discharges could be inhibited in all DNLL neurons and the inhibition often persisted for periods ranging from 10 to 50 ms beyond the duration of the tone burst that evoked it. The duration of the persistent inhibition increased with stimulus intensity. Stimulus duration had little influence on the duration of the persistent inhibition. Signals as short as 2 ms suppressed discharges for as long as 30 ms after the signal had ended. The frequency tuning of the total period of inhibition and the period of persistent inhibition were both closely matched to the tuning evoked by stimulation of the contralateral ear. Moreover, the effectiveness of complex signals for evoking persistent inhibition, such as brief FM sweeps and sinusoidally amplitude and frequency modulated signals, was comparable to that of tone bursts at the neuron's excitatory BF, so long as the complex signal contained frequencies at or around the neuron's excitatory BF. We also challenged DNLL cells with binaural paradigms. In one experiment, we presented a relatively long (40 ms) BF tone burst of fixed intensity to the contralateral ear, which evoked a sustained discharge, and a shorter, 10 ms signal of variable intensity to the ipsilateral ear. As the intensity of the 10 ms ipsilateral signal increased, it generated progressively longer periods of persistent inhibition and thus the discharges were suppressed for periods far longer than the 10 ms duration of the ipsilateral signal. With interaural time disparities, ipsilateral signals that led contralateral signals evoked a persistent inhibition that suppressed the responses to the trailing contralateral signals for periods of a least 15 ms. This suggests that an initial binaural sound that favors the ipsilateral ear should suppress the responses to trailing sounds that normally would be excitatory if they were presented alone. We hypothesize a circuit that generates the persistent inhibition and discuss how the results with binaural signals support that hypothesis.

Effects of interaural intensity and time disparity on transient evoked otoacoustic emissions

Monaural and binaural 11/s, 65 dB pe SPL clicks with interaural time and intensity disparities known to affect central auditory processing were used to study contralateral suppression of transient evoked otoacoustic emissions (TEOAEs) in 10 subjects (20 ears). Psychophysical assessment of sound lateralization induced by the same stimuli was also conducted. TEOAEs were recorded to monaural (ipsilateral to the OAE recording probe) and to binaural clicks when clicks to the contralateral ear were synchronous and symmetrical in intensity, or, in the binaural intensity disparity conditions, synchronous but 10 dB higher or 10 dB lower in the ear contralateral to the OAE recording probe. When interaural time disparities were studied, the clicks to the contralateral ear were of the same intensity throughout, but 400 μs earlier or 400 μs later than to the ear with the probe. The TEOAE components at 13–15.8 ms showed suppression, relative to monaural responses, under all binaural conditions. This contralateral suppression did not correlate with the psychophysical findings. Suppression effects were more pronounced with binaural disparity than with binaurally symmetrical clicks. Thus, although contralateral click intensity was the same with time disparities, suppression was paradoxically enhanced compared to the binaurally symmetrical stimulation. To explain these results we propose that two factors are involved in TEOAE suppression with binaural clicks: (1) contralateral intensity and (2) interaural disparity (time or intensity). The latency of the suppressions observed, the effect of interaural disparity on these suppressions, coupled with the anatomical origin of the crossed efferent fibers and the disparity sensitivity of the superior olivary complex (SOC), all suggest SOC involvement in these TEOAE suppressions.

‘Odddball’ event-related potential during REM and NREM sleep

Objectives and mehtods: Four sets of measurements were obtained from 11 patients (44–80 years old) with small, localized pontine lesions due to vascular disease: (1) Monaural auditory brain-stem evoked potentials (ABEPs; peaks I to VI); (2) Binaural ABEPs processed for their binaural interaction components (BICs) in the latency range of peaks IV to VI; (3) magnetic resonance imaging (MRI) of the brain-stem; and (4) psychoacoustics of interaural time disparity measures of binaural localization. ABEPs and BICs were analyzed for peak latencies and interpeak latency differences. Three-channel Lissajous' trajectories (3-CLTs) were derived for ABEPs and BICs and the latencies and orientations of the equivalent dipoles of ABEP and BICs were inferred from them.Results: Intercomponent latency measures of monaurally evoked ABEPs were abnormal in only 3 of the 11 patients. Consistent correlations between sites of lesion and neurophysiological abnormality were obtained in 9 of the 11 patients using 3-CLT measures of BICs. Six of the 11 patients had absence of one or more BIC components. Seven of the 11 had BICs orientation abnormality and 3 had latency abnormalities. Trapezoid body (TB) lesions (6 patients) were associated with an absent (two patients with ventral-caudal lesions) or abnormal (one patient with ventral-rostral lesions) dipole orientation of the first component (at the time of ABEPs IV), and sparing of this component with midline ventral TB lesions (two patients). A deviant orientation of the second BICs component (at the time of ABEPs V) was observed with ventral TB lesions. Psychoacoustic lateralization in these patients was biased toward the center. Rostral lateral lemniscus (LL) lesions (3 patients) were associated with absent (one patient) or abnormal (two patients) orientation of the third BICs component (at the time of ABEPs VI); and a side-biased lateralization with behavioral testing.Conclusions: These results indicate that: (1) the BICs component occurring at the time of ABEPs peak IV is dependent on ventral-caudal TB integrity; (2) the ventral TB contributes to the BICs component at the time of ABEPs peak V; and (3) the rostral LL is a contributing generator of the BICs component occurring at the time of ABEP peak VI.

1810 Effect of interaural time difference on neural activity patterns of the guinea pig auditory cortex observed by optical recording

Long-latency responses to bidirectional reversals in the laterality of a sound image were recorded from vertex with left and right earlobe references simultaneously. Laterality reversal stimuli with regular ISIs of 3 sec were obtained by alternating between +1 msec and −1 msec the interaural delay (ID) of dichotically presented 70 c/sec continous click trains. Monaural presentation of this temporal modulation did not evoke any potential or any sensation whereas, when dichotically presented, the same ID modulation was perceived as laterality reversals. Evoked potentials from 9 right-handed volunteers with normal hearing were evaluated. The recorded laterality reversal auditory evoked potentials (LRAEPs) consisted, characteristically, of a negative and a positive wave with average peak latencies of 160 msec and 260 msec, respectively. Statistical analysis indicated significant dependence of the response parameters on the direction of the sound image shifts, but the laterality of the earlobe reference did not prove to be important. The appropriateness of the stimulus parameters employed was discussed from the viewpoint of dynamic systems analysis. By contrasting with the monaural auditory off-responses recorded from the same subjects, it was demonstrated that the laterality reversal responses we obtained did not seem to be non-specific ‘rest-to-motion’ responses to any change in the acoustic background. Similarities and differences between these laterality reversal responses and various ERP components in the same latency range were also discussed. It is concluded that the LRAEPs introduced in this communication can be used in assessing the functional integrity and studying the dynamics of the neurophysiological system that accomplishes sound lateralization using cues of interaural time disparity (ITD).

Binaural masking and sensitivity to interaural delay in the inferior colliculus.

The binaural masking level difference (BMLD) is a psychophysical effect whereby signals masked by a noise at one ear become unmasked by sounds reaching the other. BMLD effects are largest at low frequencies where they depend on signal phase, suggesting that part of the physiological mechanism responsible for the BMLD resides in cells that are sensitive to interaural time disparities. We have investigated a physiological basis for unmasking in the responses of delay-sensitive cells in the central nucleus of the inferior colliculus in anaesthetized guinea pigs. The masking effects of a binaurally presented noise, as a function of the masker delay, were quantified by measuring the number of discharges synchronized to the signal, and by measuring the masked threshold. The noise level for masking was lowest at the best delay for the noise. The mean magnitude of the unmasking across our neural population was similar to the human psychophysical BMLD under the same signal and masker conditions.

Onset coding of spatial information in trains of identical transients

In a study of the effects of onsets in lateralizing bursts of noise, Tobias and Schubert [J. Acoust. Soc. Am. 31, 1595–1605 (1959)] demonstrated that, depending on the duration of the ongoing noise burst, it takes anywhere between 4 to 35 times as much onset interaural time disparity to offset ongoing disparities. They, therefore, demonstrated that onsets do not necessarily provide the major cue to auditory localization. In the present study, onset and ongoing information in a lateralization paradigm is set in spatial conflict to determine the amount of onset interaural difference of time (IDT) necessary to offset ongoing IDTs of transient trains of various durations. The major difference between this study and the Tobias and Schubert study was that this study used identical transients [e.g., interclick interval (ICI) = 1.5 to 2 ms corresponding to a 500‐ to 750‐Hz tone] and not noise bursts. Unlike the Tobias and Schubert study, it was found in this study that a single dichotic click could easily lateralize a contralateral transient train of up to 200 clicks; producing the sensation of a tone lateralized to the position of the initial event. Various ICIs and ongoing durations were tested. In such a context, the precedence effect, the Franssen illusion, and the formation of auditory objects will be discussed. [Work supported by NSE and NIH.]

Human laterality reversal auditory evoked potentials: stimulation by reversing the interaural delay of dichotically presented continuous click trains

Long-latency responses to bidirectional reversals in the laterality of a sound image were recorded from vertex with left and right earlobe references simultaneously. Laterality reversal stimuli with regular ISIs of 3 sec were obtained by alternating between +1 msec and -1 msec the interaural delay (ID) of dichotically presented 70 c/sec continuous click trains. Monaural presentation of this temporal modulation did not evoke any potential or any sensation whereas, when dichotically presented, the same ID modulation was perceived as laterality reversals. Evoked potentials from 9 right-handed volunteers with normal hearing were evaluated. The recorded laterality reversal auditory evoked potentials (LRAEPs) consisted, characteristically, of a negative and a positive wave with average peak latencies of 160 msec and 260 msec, respectively. Statistical analysis indicated significant dependence of the response parameters on the direction of the sound image shifts, but the laterality of the earlobe reference did not prove to be important. The appropriateness of the stimulus parameters employed was discussed from the viewpoint of dynamic systems analysis. By contrasting with the monaural auditory off-responses recorded from the same subjects, it was demonstrated that the laterality reversal responses we obtained did not seem to be non-specific 'rest-to-motion' responses to any change in the acoustic background. Similarities and differences between these laterality reversal responses and various ERP components in the same latency range were also discussed. It is concluded that the LRAEPs introduced in this communication can be used in assessing the functional integrity and studying the dynamics of the neurophysiological system that accomplishes sound lateralization using cues of interaural time disparity (ITD).