Auditory cortical responses to abrupt lateralization shifts do not reflect the activity of hemifield-specific units involved in opponent coding of auditory space.

Recent studies show that the classical model based on axonal delay-lines may not explain interaural time difference (ITD) based spatial coding in humans. Instead, a population-code model called "opponent channels model" (OCM) has been suggested. This model comprises two competing channels respectively for the two auditory hemifields, each with a sigmoidal tuning curve. Event-related potentials (ERPs) to ITD-changes are used in some studies to test the predictions of this model by considering the sounds before and after the change as adaptor and probe stimuli, respectively. It is assumed in these studies that the former stimulus causes adaptation of the neurons selective to its side, and that the ERP N1-P2 response to the ITD-change is the specific response of the neurons with selectivity to the side of probe sound. However, these ERP components are known as a global, non-specific acoustic change complex of cortical origin evoked by any change in the auditory environment. It probably does not genuinely reflect the activity of some stimulus-specific neuronal units that have escaped the refractory effect of the preceding adaptor, which means a violation of the crucial assumption in an adaptor-probe paradigm. To assess this viewpoint, we conducted two experiments. In the first one, we recorded ERPs to abrupt lateralization shifts of click trains having various pre- and post-shift ITDs within the physiological range of -600µs to +600µs. Magnitudes of the ERP components P1, N1, and P2 to these ITD-shifts did not comply with the additive behavior of partial probe responses presumed for an adaptor-probe paradigm, casting doubt on the accuracy of testing sensory coding models by using ERPs to abrupt lateralization changes. Findings of the second experiment, involving ERPs to conjoint outwards/transverse shift stimuli also supported this conclusion.

Estimating and minimizing movement artifacts in surface electromyogram.

While recording surface electromyography [sEMG], it is possible to record the electrical activities coming from the muscles and transients in the half-cell potential at the electrode-electrolyte interface due to micromovements of the electrode-skin interface. Separating the two sources of electrical activity usually fails due to the overlapping frequency characteristics of the signals. This paper aims to develop a method that detects movement artifacts and suggests a minimization technique. Towards that aim, we first estimated the frequency characteristics of movement artifacts under various static and dynamic experimental conditions. We found that the extent of the movement artifact depended on the nature of the movement and varied from person to person. Our study's highest movement artifact frequency for the stand position was 10 Hz, tiptoe 22, walk 32, run 23, jump from box 41, and jump up and down 40 Hz. Secondly, using a 40 Hz highpass filter, we cut out most of the frequencies belonging to the movement artifacts. Finally, we checked whether the latencies and amplitudes of reflex and direct muscle responses were still observed in the highpass-filtered sEMG. We showed that the 40 Hz highpass filter did not significantly alter reflex and direct muscle variables. Therefore, we recommend that researchers who use sEMG under similar conditions employ the recommended level of highpass filtering to reduce movement artifacts from their records. However, suppose different movement conditions are used. In that case, it is best to estimate the frequency characteristics of the movement artifact before applying any highpass filtering to minimize movement artifacts and their harmonics from sEMG.

Effects of aging on event-related potentials to single-cycle binaural beats and diotic amplitude modulation of a tone.

Aim of the study is to determine whether the auditory processing of temporal fine structure (TFS) is affected with normal aging, even in the presence of normal audiometric hearing and fine cognitive state; and, if it is, to see whether a comparable effect is also observed in the processing of a diotic change in sound envelope. The event-related potentials (ERPs) to binaural beats (BBs), which are the responses of the binaural mechanisms processing TFS of a sound, and the ERPs to diotic amplitude modulation (AM) stimuli, which are the responses of the monaural mechanisms processing the changes in its envelope, were recorded from thirteen young university students and ten senior but active university professors, all with normal hearing in low frequencies. To obtain directly the specific BB responses without confounding monaural frequency change-evoked responses, we used single-cycle BB stimuli with temporary sub-threshold frequency shifts. BBs of a 250-Hz tone and diotic AM of the same tone with similar perceptual salience were presented with 2-second stimulus onset asynchrony. The N1 components of the ERPs to both stimuli displayed notable age-dependent changes in their scalp topography and significant amplitude reduction and latency prolongation in the elderly. These amplitude and latency changes were at similar rates for the two stimulus types, implying that the auditory TFS and envelope processing mechanisms are proportionally affected by physiological aging. These results may serve as control data in future studies investigating the effect of aging-associated cognitive pathologies on auditory TFS processing.

Event-related potentials to single-cycle binaural beats of a pure tone, a click train, and a noise.

There are only few electrophysiological studies on a phenomenon called "binaural beats" (BBs), which is experienced when two tones with frequencies close to each other are dichotically presented to the ears. And, there is no study in which the electrical responses of the brain to BBs of complex sounds are recorded and analyzed. Owing to a recent method based on single-cycle BB stimulation with sub-threshold temporary monaural frequency shifts, we could record the event-related potentials (ERPs) to BBs of a 250-Hz tone as well as those to the BBs of a 250/s click train and to the BBs of a recurrent 4-ms Gaussian noise. Although fundamental components of the click train and noise stimuli were lower in intensity than the tonal stimuli in our experiments, the N1 responses to the BBs of the former two wide-spectrum sounds were recorded with significantly larger amplitudes and shorter latencies than those to the BBs of a tone, suggesting an across-frequency integration of directional information. During a BB cycle of a complex sound, the interaural time differences (ITDs) of the spectral components are all equal to each other at any time; whereas their interaural phase differences (IPDs) are all different. The ITD rather than the IPD should, therefore, be the cue that is relied upon by the binaural mechanism coding the perceived lateral shifts of the sound caused by BBs. This is in line with across-frequency models of human auditory lateralization based on a common ITD, fulfilling a straightness criterion.

Facial muscle activity contaminates EEG signal at rest: evidence from frontalis and temporalis motor units.

OBJECTIVE: In order to reach electroencephalography (EEG) electrodes on the scalp, synchronized activity of neurons needs to pass thorough several tissue layers, including the skull and muscles covering the scalp. The contamination of EEG signal by temporalis and frontalis muscles has been well documented for voluntary muscle contraction even at low contraction levels. The extent of myogenic contamination during postural and/or rest activity of the temporalis and frontalis remains an impediment for EEG research. APPROACH: In this study, we first aimed to observe involuntary, continuous motor unit activity of the frontalis muscle at rest and evaluate motor unit level frontalis interference on the EEG electrodes. Second, we compared motor unit interference from the frontalis before and after artefact pruning via an independent component analysis (ICA) algorithm. MAIN RESULTS: We demonstrated that motor unit activity of the frontalis muscle produces interference potentials on the frontal electrodes at rest and the interference was significantly reduced after ICA on the frontal electrodes, but not completely eliminated. Likewise, the temporalis interference at rest was significantly smaller after ICA on the fronto-temporal electrodes, but not completely removed. SIGNIFICANCE: We documented the existence of resting involuntary activity of the temporalis and frontalis muscles underneath EEG electrodes and the removal of the EEG signal from their contiguous interference is not possible even after the use of ICA technology. We recommend that EEG researchers readdress the definition of 'rest' for EEG recordings and the ICA experts should extend their electromyography removal strategies to motor unit level interference.

Event-related potentials to single-cycle binaural beats and diotic amplitude modulation of a tone.

When two tones with slightly different frequencies are dichotically presented, binaural beats (BBs) are experienced. BBs resulting from the cycling change in interaural phase difference elicit electroencephalographic responses. Because they repeat at short periods, allowing poor recovery of the cortical responses, these steady-state responses have small amplitudes, and their various wave components intermingle and might mask each other. Using single-cycle BBs separated by relatively long inter-onset intervals would be a solution, but introducing a transient interaural frequency shift requires response subtraction which may not be acceptable for non-additive brain responses. The proposed stimulation method employs transient and monaurally subthreshold frequency shifts in opposite directions in the two ears to produce single-cycle BBs of a 250 Hz tone. These shifts are perceived as distinct BBs when presented dichotically, but remain subthreshold when presented monotically. Therefore, no frequency-shift response is elicited, and the specific BB response is obtained with no need for waveform subtraction. We recorded from 19 normal hearing participants the event-related potentials (ERPs) to single-cycle BBs and also to temporary diotic amplitude modulation (AM) with matched perceptual salience. The ERPs to single-cycle BBs presented at 2 s inter-onset intervals had N1-P2 responses with up to seven times larger amplitudes than the conventional steady-state BB responses in the literature. Significant differences were found between the scalp potential distributions of the N1 responses to BB and AM stimuli, suggesting that the cortical sites, where envelope-based level processing and temporal fine structure-based spatial processing of the stimulus take place, are not totally overlapped.

Pre-attentive Mismatch Response and Involuntary Attention Switching to a Deviance in an Earlier-Than-Usual Auditory Stimulus: An ERP Study.

An acoustic stimulus elicits an electroencephalographic response called auditory event-related potential (ERP). When some members of a stream of standard auditory stimuli are replaced randomly by a deviant stimulus and this stream is presented to a subject who ignores the stimuli, two different ERPs to deviant and standard stimuli are recorded. If the ERP to standard stimuli is subtracted from the ERP to deviant stimuli, the difference potential (DP) waveform typically exhibits a series of negative-positive-negative deflections called mismatch negativity (MMN), P3a, and reorienting negativity (RON), which are associated with pre-attentive change detection, involuntary attention switching, and reorienting of attention, respectively. The aim of the present study was to investigate how these pre-attentive processes are affected if the change occurs earlier than its usual timing implied by isochronous standard stimuli. In the MMN paradigm employed, 15% of the standards were randomly replaced by deviant stimuli which differed either in their pitch, their earlier onset time, or in both. Event-related responses to these three deviants [timely pitch change (RTP), earlier onset (REO), earlier pitch change (REP)] and to standards (RS) were recorded from 10 reading subjects. To maintain identical stimulation histories for the responses subtracted from each other, "deviant-standard" difference potentials (DP) for "timely" and "early" pitch deviances were derived as follows: DPTP = RTP - RS and DPEP = REP - REO. Interestingly, the MMN components of the DPs to timely and early pitch deviances had similar amplitudes, indicating that regularity of stimulus timing does not provide any benefit for the pre-attentive auditory change detection mechanism. However, different scalp current density (SCD) dynamics of the MMN/P3a complexes, elicited by timely and early pitch deviances, suggested that an auditory change in a stimulus occurring earlier-than-usual initiates a faster and more effective call-for-attention and causes stronger attention switching than a timely change. SCD results also indicated that the temporal, frontal, and parietal MMN components are simultaneously present rather than emerging sequentially in time, supporting the MMN models based on parallel deviance processing in the respective cortices. Similarity of the RONs to timely and early pitch deviances indicated that reorienting of attention is of the same strength in two cases.

EEG-like signals can be synthesized from surface representations of single motor units of facial muscles.

Electrodes for recording electroencephalogram (EEG) are placed on or around cranial muscles; hence, their electrical activity may contaminate the EEG signal even at rest conditions. Due to its role in maintaining mandibular posture, tonic activity of temporalis muscle interferes with the EEG signal particularly at fronto-temporal locations at single motor unit (SMU) level. By obtaining surface representation of a motor unit, we can evaluate its interference in EEG and if we could sum surface representations of several tonically active motor units, we could estimate the overall myogenic contamination in EEG. Therefore, in this study, we followed re-composition (RC) approach and generated EEG-like artefact model using surface representations of single motor units (RC). Furthermore, we compared signal characteristics of RC signals with simultaneously recorded EEG signal at different locations in terms of power spectral density and coherence. First, we found that RC signal represented the power spectral distribution of an EMG signal. Second, RC signal reflected the discharge rate of a SMU giving the greatest surface representation amplitude and strongest interference appeared as distinguishable frequency peak on RC power spectra. Moreover, for strong interferences, RC also contaminated the EEG at F7 and other EEG electrodes. These findings are important to illustrate the susceptibility of EEG signal to myogenic artefacts even at rest and the research using EEG coherence comparisons should consider muscle activity while drawing conclusions about neuronal interactions and oscillations.

Interference of tonic muscle activity on the EEG: a single motor unit study.

The electrical activity of muscles can interfere with the electroencephalogram (EEG) signal considering the anatomical locations of facial or masticatory muscles surrounding the skull. In this study, we evaluated the possible interference of the resting activity of the temporalis muscle on the EEG under conventional EEG recording conditions. In 9 healthy adults EEG activity from 19 scalp locations and single motor unit (SMU) activity from anterior temporalis muscle were recorded in three relaxed conditions; eyes open, eyes closed, jaw dropped. The EEG signal was spike triggered averaged (STA) using the action potentials of SMUs as triggers to evaluate their reflections at various EEG recording sites. Resting temporalis SMU activity generated prominent reflections with different amplitudes, reaching maxima in the proximity of the recorded SMU. Interference was also notable at the scalp sites that are relatively far from the recorded SMU and even at the contralateral locations. Considering the great number of SMUs in the head and neck muscles, prominent contamination from the activity of only a single MU should indicate the susceptibility of EEG to muscle activity artifacts even under the rest conditions. This study emphasizes the need for efficient artifact evaluation methods which can handle muscle interferences.

Electrophysiological assessment of the effects of obstructive sleep apnea on cognition.

We used electrophysiological measures to investigate the effects of obstructive sleep apnea on attention, learning, and memory. Thirty subjects (OSA group, n = 15, control group n = 15) participated in n-back tests, accompanied by P300 recordings, to investigate working memory and attention. The mirror-drawing test was used to study procedural memory, and the trail-making test (TMT) was used to evaluate divided attention and executive function. No significant group difference in reaction time was found in the 0-back and 1-back tests. In the 2-back test, reaction times of patients were longer than those of the control group. No P300 wave was obtained in the OSA group in any (0-, 1-, or 2-back) n-back test. In contrast, in the control group, significant P300 waves were recorded except for the 2-back test. The mirror-drawing scores were unaffected by sleep apnea. There was no difference between groups in the TMT-A test on any of the trials. Although no group difference was found in the first or second trials of the TMT-B test, OSA patients were less successful in learning on the third trial. According to our study results, OSA affects attention and executive function adversely however, we could not detect a significant effect on working or procedural memory.

Significant variations in Weber fraction for changes in inter-onset interval of a click train over the range of intervals between 5 and 300 ms.

It is a common psychophysical experience that a train of clicks faster than ca. 30/s is heard as one steady sound, whereas temporal patterns occurring on a slower time scale are perceptually resolved as individual auditory events. This phenomenon suggests the existence of two different neural mechanisms for processing of auditory sequences with fast and slow repetition rates. To test this hypothesis we used Weber's law, which is known to be valid for perception of time intervals. Discrimination thresholds and Weber fractions (WFs) for 12 base inter-click intervals (ICIs) between 5 and 300 ms were measured from 10 normal hearing subjects by using an "up-down staircase" algorithm. The mean WF, which is supposed to be constant for any perceptual mechanism according to Weber's law, displayed significant variation with click rate. WFs decreased sharply from an average value of around 5% at repetition rates below 20 Hz to about 0.5% at rates above 67 Hz. Parallel to this steep transition, subjects reported that at rates below 20 Hz they perceived periodicity as a fast tapping rhythm, whereas at rates above 50 Hz the perceived quality was a pitch. Such a dramatic change in WF indicated the existence of two separate mechanisms for processing the click rate for long and short ICIs, based on temporal and spectral features, respectively. A range of rates between 20 and 33 Hz, in which the rate discrimination threshold was maximum, appears to be a region where both of the presumed time and pitch mechanisms are relatively insensitive to rate alterations. Based on this finding, we speculate that the interval-based perception mechanism ceases to function at around 20 Hz and the spectrum-based mechanism takes over at around 33 Hz; leaving a transitional gap in between, where neither of the two mechanisms is as sensitive. Another notable finding was a significant drop in WF for ICI = 100 ms, suggesting a connection of time perception to the electroencephalography alpha rhythm.

Event-related potentials to changes of rhythmic unit: differences between musicians and nonmusicians.

We investigated whether the expected differences between musicians and nonmusicians in their ability to detect a rhythm change were reflected in their event-related potentials (ERPs) and, if reflected, how these ERP differences associated with behavioral indices. Stimuli were three consecutive and equally spaced drum beats followed by a rest period to form a rhythmic unit (RU). By using three different inter-beat periods, three RUs were produced. Combinations of these RUs served as the "target/standard" pairs of an oddball sequence. In four different experiments, we tried two RU-change types each with two levels of detection difficulty. ERPs were recorded from the F3, Fz, F4, Cz and Pz scalp sites of 12 musicians and 12 nonmusicians. RT, hit and false-alarm rates were also measured. The data have shown with high statistical confidence that, associated with the musicians' better detection performance and shorter RTs, their ERP P3 to rhythm changes peaked significantly earlier and was significantly larger compared to nonmusicians. Intergroup ERP differences allowed above 90% correct classification. This study has also showed that not only violations of relatively complex musical regularities, but very simple rhythmic unit alterations could lead to significant P3 differences between musicians and nonmusicians. The high accuracy of the musician/nonmusician classification based only on their P3 data strongly supported the hypothesis that sensory and/or cognitive advantage of musicians in detecting rhythm changes does reflect in their P3.

Lateralisation of sound in temporal-lobe epilepsy: comparison between pre- and postoperative performances and ERPs.

OBJECTIVE: Our aim was to investigate if spatial hearing is impaired in mesial temporal lobe epilepsy and temporal lobectomy has an effect on this function. METHODS: Thirteen patients with mesial temporal lobe epilepsy (TLE) due to sclerosis in their left (n=6) or right (n=7) hippocampus were studied. Their sound lateralisation performance indexed by d' was tested against that of a group of normal subjects (n=13). Patients' ERPs to lateralisation shifts induced by interaural disparities of intensity (IID) and time (ITD) were also recorded. Eight of the patients were re-tested after they underwent anterior temporal lobectomy, which involved the resection/removal of medial structures including amygdala, hippocampus and parahippocampal gyrus. RESULTS: The sound-lateralisation performance of the TLE patients was significantly lower than normal subjects, and this disadvantage of the patients was specific to IID-based lateralisation. Amplitudes of their N1 and P2 responses to laterally shifting sounds were much lower than those reported previously for normal subjects. Lobectomy did not have a statistically significant effect on patients' sound-lateralisation performance nor on the amplitude of their auditory directional ERPs. CONCLUSIONS: The results show that especially the IID-based sound-lateralisation performance is impaired in TLE patients and that lobectomy should not cause any further deterioration. SIGNIFICANCE: This study suggests that a test for assessing the ability of sound lateralisation based on each of the IID and ITD cues should be included in the evaluation of TLE patients.

Assessment of the role of the cochlear latency effect in lateralization of click sounds in humans.

Interaural time and intensity disparities (ITD and IID) are the two cues to sound lateralization. "Time-only" hypothesis claims that an IID is first converted to an interaural afferent delay (Delta t), and is then processed by the central ITD mechanism, rendering a separate IID processor unnecessary. We tested this hypothesis by assessing the contribution of the cochlear latency effect to the psychophysical ITD/IID trading ratio. Auditory brainstem responses (ABRs) were used to measure the interaural afferent delays (Delta ts) that developed with a 20/sec dichotic click train used in the trading experiment. Except for small IIDs at low loudness levels, the physiological Delta t delay produced by an IID was significantly smaller than the ITD psychophysically traded for the same IID. We concluded that the cochlear latency effect alone cannot explain the psychophysical ITD/IID trading ratios and a separate IID mechanism must be involved.

Refractoriness, habituation, and mismatch effects in the auditory event-related potential to alternating stimuli.

Amplitude enhancement in the N1 component of auditory event-related potentials (ERPs) to alternately presented sounds has been referred as a typical example for the effect of release from neural refractoriness. We tested this hypothesis to see whether some other effects also contribute to this phenomenon. Two tones of different frequencies were presented singly or in pairs, and ERPs were recorded using monotonous (mnt) and alternating (alt) sequences of these stimuli. Comparison of the 'alt-mnt' difference waveforms recorded with single and paired stimuli supported the refractoriness hypothesis. A mismatch negativity-like wave, however, was also observed, questioning the constraint of 'at least two consecutive standards before deviant' presumed in most mismatch negativity studies. This paradigm made it possible to delineate the ERP components related to refractoriness and mismatch detection processes.

Changes in the alpha and beta amplitudes of the central EEG during the onset, continuation, and offset of long-duration repetitive hand movements.

Electroencephalographic alpha and beta activities recorded from central electrodes are known to display movement-related suppression or enhancement. We investigated whether the suppression that is known to occur during the onset of a single movement would persist or otherwise habituate when the movement is continuously repeated for a long period of time. Fourteen subjects took part in the experiments. They performed repetitive simultaneous extension-flexions of the fingers II-V in one hand, continuously for a period of at least 30 s. They then stopped this self-paced movement and rested for at least 30 s. Bipolar recording was made from C3-Cz and C4-Cz. Patterns of amplitude changes in the alpha and beta bands were calculated against a resting baseline. Following a bilateral alpha and beta suppression at the movement onset, alpha amplitude gradually but not fully recovered towards the baseline during the 30 s post-onset. Habituation of afferences and transfer of the cortical function were discussed as the two alternative explanations for this gradual recovery. Beta amplitude, however, displayed no recovery as long as the movement continued. Considering the relatively rapid beta recovery reported for sustained movements, this finding demonstrated that the sustained and continuous movements are conducted through quite different processes. A transient contralateral beta rebound was observed only after the end of the long movement period, strengthening the viewpoint that links the beta rebound with the closure of the cortical processes running throughout a motor sequence. Modulation of the beta amplitude, rather than the changes in alpha amplitude, appeared to be more closely correlated with the execution of a continuous movement.

The 'Franssen' illusion for short duration tones is preattentive: a study using mismatch negativity.

When a tone burst is divided into two parts, an onset transient and a sustained tone smoothly fading on, and these parts are delivered to two stereophonically located loudspeakers in a room, a listener gains the impression that the whole sound is coming from the loudspeaker that actually emits merely the transient. Due to this auditory illusion known as the 'Franssen effect' (FE), the physical and the perceived lateralizations of the sustained sound become different. A two-block mismatch negativity (MMN) paradigm was used to investigate the stage of auditory processing at which this segregation would take place. In one block, standard stimuli were 100 ms, 1 kHz tone bursts emitted by one of the loudspeakers, and deviant stimuli were their split version, with the sustained part switched to the other loudspeaker. In the other block, the roles of the two stimuli were swapped. A room acoustics software was used for generating the signals to a headphone. The responses recorded from 10 subjects displayed no MMN, although the same stimuli but without the transients evoked prominent MMNs. This indicated that the mechanism underlying this illusion modifies the neural representation of the stimulus with FE in such a way that it becomes similar to that of the stimulus without FE before reaching the input of the preattentive mechanism indexed by the MMN. Considering the possible relationship of this illusion to the precedence effect and also the relevant electrophysiological findings in the literature, we conclude that the primary auditory cortex is the most plausible site of the mechanism leading to the FE.

Interaural delay-dependent changes in the binaural interaction component of the guinea pig brainstem responses.

Auditory brainstem responses to monaural and binaural clicks with 23 different interaural time differences (ITDs) were recorded from ten guinea pigs without anesthesia. Binaural interaction component was obtained by subtracting the sum of the appropriately time-shifted left and right monaural responses from the binaural one. With increasing ITD, the most prominent peak of the binaural difference potential so obtained shifted to longer latencies and its amplitude gradually decreased. The way these changes depended on binaural delay was basically similar to that previously observed in a cat study [P. Ungan, S. Yagcioglu, B. Ozmen. Interaural delay-dependent changes in the binaural difference potential in cat auditory brainstem response: implications about the origin of the binaural interaction component. Hear. Res. 106 (1997) 66-82]. The data were successfully simulated by the model suggested in that report. We therefore concluded that the same model, which was based on the difference between the mean onset latencies of the ipsilateral excitation and contralateral inhibition in a typical neuron in the lateral superior olive, their standard deviations, and the duration of the contralateral inhibition, should also be valid for the binaural interaction in the guinea pig brainstem. The results, which were discussed in connection with sound lateralization models, supported a model based on population coding, where the lateral position of a sound source is coded by the ratio of the discharge intensity in the left and right lateral superior olives, rather than the models based on coincidence detection.

Dynamics of the contralateral white noise-induced enhancement in the guinea pig's middle latency response.

The peak-to-peak amplitude of temporal middle latency response (MLR) of the guinea pig, evoked by a click in the contralateral ear, according to the recording side, is increased with the presence of continuous white noise (CWN) in the ipsilateral ear and this specialty is defined as the white noise enhancement (WNE). This phenomenon is evaluated as an interesting electrophysiological finding from the viewpoint of binaural interaction and in this study, its dynamic specifications were investigated. After the beginning of ipsilateral CWN, significant WNE was observed at 275th ms and it reached to a maximum, with an increase more than 40%, at 350th ms. After a habituation occurred, WNE reached to 20% on the 4th second by gradually decreasing and came to a steady state. In the time window between 2 and 5 ms after CWN started, a surprising amplitude decrease is observed. Therefore, CWN causes an effect, like a click, in the short-term and this on-response type effect originates from low level binaural centers, which decreases the MLR amplitude. However, the same CWN increases the MLR amplitude (WNE) by the effects over the high level binaural centers in the succeeding period, by its continuous characteristic.

Origin of the binaural interaction component in wave P4 of the short-latency auditory evoked potentials in the cat: evaluation of serial depth recordings from the brainstem.

There is no general agreement on the origin of the binaural interaction (BI) component in auditory brainstem responses (ABRs). To study this issue the ABRs to monaural and binaural clicks with various interaural time differences (ITDs) were simultaneously recorded from the vertex and from a recording electrode aiming at the superior olive (SO) in cats. Electrode path was along the fibers of the lateral lemniscus (LL). Binaural difference potentials (BDPs), which were computed by subtracting the sum of the two monaural responses from the binaural response, were obtained at systematic depths and across a range of ITD values. It was observed that only a specific BDP deflection recorded at the level at which lemniscal fibers terminate in the nuclei of LL coincided in time with the most prominent BDP in the cat's vertex-recorded ABRs, the BDP in their wave P4. As ITD was increased, the latency shifts and amplitude decrements of the scalp-recorded far-field BDP wave exactly followed those recorded at this lemniscal near-field BDP locus. The data support our hypothesis that the BI component in wave P4 results from a binaural reduction in dischargings of axons ascending in the LL, with this reduction due to contralateral inhibition of the discharge activity of the inhibitory-excitatory units in the lateral nucleus of the SO. Furthermore, at the level of the SO, the BDP in the responses to contra-leading binaural clicks always had larger magnitudes than those evoked by ipsi-leading ones. This bilateral asymmetry is consistent with the view that the BDP in scalp-recorded ABRs is related to the function of sound lateralization.