Binaural Difference Potential Research Articles

Zur Beziehung zwischen den binauralen Differenzpotentialen und dem Richtungshörvermögen

Central auditory processing disorders (CAPD) are associated with reduced discriminatory abilities of the auditory system. One of these abilities is directional hearing, which is based on the evaluation of interaural signal differences. Since these differences affect also the binaural difference potentials (BDP), these derived evoked potentials could be suitable for the objective detection of disabilities in acoustic localization. Auditory brainstem responses (ABR) evoked by monaural and binaural stimulation were recorded in 32 adult normal-hearing volunteers at stimulus levels between 10 and 80 dB nHL and the BDP were derived. Additionally, a test of directional hearing was performed and evaluated. BDP are detectable at all stimulus levels without difficulties, with acceptable examination times and with satisfactory reliability in a routine clinical environment. The electrodes can be positioned at the mastoid (A1 and A2) and vertex (Cz) as for conventional ABR recording. Normal ranges are given for latency and amplitude as well as for their lateral differences. The spectrum in directional hearing ability as covered by normal-hearing subjects is not mirrored in the parameters of their BDP. Nevertheless, it cannot be ruled out that a real impairment of directional hearing goes along with significantly altered BDP.

The influence of externalization and spatial cues on the generation of auditory brainstem responses and middle latency responses

The effect of externalization and spatial cues on the generation of auditory brainstem responses (ABRs) and middle latency responses (MLRs) was investigated in this study. Most previous evoked potential studies used click stimuli with variations of interaural time (ITDs) and interaural level differences (ILDs) which merely led to a lateralization of sound inside the subject’s head. In contrast, in the present study potentials were elicited by a virtual acoustics stimulus paradigm with ‘natural’ spatial cues and compared to responses to a diotic, non-externalized reference stimulus. Spatial sound directions were situated on the horizontal plane (corresponding to variations in ITD, ILD, and spectral cues) or the midsagittal plane (variation of spectral cues only). An optimized chirp was used which had proven to be advantageous over the click since it compensates for basilar membrane dispersion. ABRs and MLRs were recorded from 32 scalp electrodes and both binaural potentials (B) and binaural difference potentials (BD, i.e., the difference between binaural and summed monaural responses) were investigated. The amplitudes of B and BD to spatial stimuli were not higher than those to the diotic reference. ABR amplitudes decreased and latencies increased with increasing laterality of the sound source. A rotating dipole source exhibited characteristic patterns in dependence on the stimulus laterality. For the MLR data, stimulus laterality was reflected in the latency of component N a. In addition, dipole source analysis revealed a systematic magnitude increase for the dipole contralateral to the azimuthal position of the sound source. For the variation of elevation, the right dipole source showed a stronger activation for stimuli away from the horizontal plane. The results indicate that at the level of the brainstem and primary auditory cortex binaural interaction is mostly affected by interaural cues (ITD, ILD). Potentials evoked by stimuli with natural combinations of ITD, ILD, and spectral cues were not larger than those elicited by diotic chirps.

Interaural delay-dependent changes in the binaural difference potential of the human auditory brain stem response

Binaural difference potentials (BDs) are thought to be generated by neural units in the brain stem responding specifically to binaural stimulation. They are computed by subtracting the sum of monaural responses from the binaural response, BD = B − (L + R). BDs in dependency on the interaural time difference (ITD) have been measured and compared to the Jeffress model in a number of studies with conflicting results. The classical Jeffress model assuming binaural coincidence detector cells innervated by bilateral excitatory cells via two delay lines predicts a BD latency increase of ITD/2. A modification of the model using only a single delay line as found in birds yields a BD latency increase of ITD. The objective of this study is to measure BDs with a high signal-to-noise ratio for a large range of ITDs and to compare the data with the predictions of some models in the literature including that of Jeffress. Chirp evoked BDs were recorded for 17 ITDs in the range from 0 to 2 ms at a level of 40 dB nHL for four channels (A1, A2, PO9, PO10) from 11 normal hearing subjects. For each binaural condition 10,000 epochs were collected while 40,000 epochs were recorded for each of the two monaural conditions. Significant BD components are observed for ITDs up to 2 ms. The peak-to-peak amplitude of the first components of the BD, DP1-DN1, is monotonically decreasing with ITD. This is in contrast with click studies which reported a constant BD-amplitude for ITDs up to 1 ms. The latency of the BD-component DN1 is monotonically, but nonlinearly increasing with ITD. In the current study, DN1 latency is found to increase faster than ITD/2 but slower than ITD incompatible with either variant of the Jeffress model. To describe BD waveforms, the computational model proposed by Ungan et al. [Hearing Research 106, 66–82, 1997] using ipsilateral excitatory and contralateral inhibitory inputs to the binaural cells was implemented with only four parameters and successfully fitted to the BD data. Despite its simplicity the model predicts features which can be physiologically tested: the inhibitory input must arrive slightly before the excitatory input, and the duration of the inhibition must be considerably longer than the standard deviations of excitatory and inhibitory arrival times to the binaural cells. With these characteristics, the model can accurately describe BD amplitude and latency as a function of the ITD.

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.

Auditory evoked potentials to changes in interaural cross correlation

The objective of this study is to analyze the representation of specific binaural stimulus features in the human brain and to compare the results to psychoacoustic data. Late auditory evoked potentials to changes in the interaural cross correlation (IACC) of continuous bandpass noise (100–2000 Hz) were recorded, 1000 sweeps per stimulus condition were averaged. In experiment 1 a brief ‘‘binaural gap’’ with a gap IACC was interposed in the noise with a reference IACC. N1 and P2 amplitudes increased with increasing gap duration. For uncorrelated reference IACC (0/1/0 and 0/−1/0 conditions) objective thresholds were comparable to psychoacoustic data. For uncorrelated gap IACC (1/0/1 and −1/0/−1 conditions) objective thresholds were about four times larger than in psychoacoustic measurements. In experiment 2 reference and test interval had the same length of 800 ms and the magnitude of change in IACC was varied. Objective IACC-JNDs are larger for an uncorrelated reference than for IACCs of ±1. For the uncorrelated reference, test IACCs towards +1 elicited larger AEP components than test IACCs towards −1. In contrast to binaural difference potentials used at the brain stem level, employing changes in IACC constitutes a more direct approach to objectively investigate the binaural system in humans.

Examination of binaural signal processing in normally hearing subjects using electrophysiological and psychoacoustical measurements

At present, only a small number of validated, clinically usable methods for the assessment of binaural hearing capabilities exist. A proposed electrophysiological measure is the registration of the brainstem-based binaural difference potentials (BDP). The BDP is calculated as the difference between the binaurally evoked registration and the sum of the two monaural registrations. Detection and stability of the BDP were examined in 24 normally hearing adults within the framework of conventional registration of auditory brainstem responses. Furthermore, the influence of interaural time differences (ITD) on the BDP was determined. In addition, lateralization of the subjects was assessed using a psychoacoustical method. The components of the BDP could be detected in almost all of the subjects. Moreover, they showed sufficient test-retest reliability. The impact of ITD,which causes lateralization of the stimulus,was clearly detectable for the latencies and the amplitudes of the BDP. Binaural difference potentials, which are easily and reliably detectable reveal a relationship to the outcome of psychoacoustical assessment of lateralization and have the potential to provide a measure for binaural hearing capacity.

Comparison of binaural auditory brainstem responses and the binaural difference potential evoked by chirps and clicks

Rising chirps that compensate for the dispersion of the travelling wave on the basilar membrane evoke larger monaural brainstem responses than clicks [Dau et al., J. Acoust. Soc. Am. 107 (2000) 1530–1540]. In order to test if a similar effect applies for the early processing stages of binaural information, monaurally and binaurally evoked auditory brainstem responses were recorded for clicks and chirps for levels from 10 to 60 dB nHL in steps of 10 dB. Ten thousand sweeps were collected for every stimulus condition from 10 normal hearing subjects. Wave V amplitudes are significantly larger for chirps than for clicks for all conditions. The amplitude of the binaural difference potential, DP1–DN1, is significantly larger for chirps at the levels 30 and 40 dB nHL. Both the binaurally evoked potential and the binaural difference potential exhibit steeper growth functions for chirps than for clicks for levels up to 40 dB nHL. For higher stimulation levels the chirp responses saturate approaching the click evoked amplitude. For both stimuli the latency of DP1 is shorter than the latency of the binaural wave V, which in turn is shorter than the latency of DN1. The amplitude ratio of the binaural difference potential to the binaural response is independent of stimulus level for clicks and chirps. A possible interpretation is that with click stimulation predominantly binaural interaction from high frequency regions is seen which is compatible with a processing by contralateral inhibitory and ipsilateral excitatory (IE) cells. Contributions from low frequencies are negligible since the responses from low frequencies are not synchronized for clicks. The improved synchronization at lower frequencies using chirp stimuli yields contributions from both low and high frequency neurons enlarging the amplitudes of the binaural responses as well as the binaural difference potential. Since the constant amplitude ratio of the binaural difference potential to the binaural response makes contralateral and ipsilateral excitatory interaction improbable, binaural interaction at low frequencies is presumably also of the IE type. Another conclusion of this study is that the chirp stimuli employed here are better suited for auditory brainstem responses and binaural difference potentials than click stimuli since they exhibit higher amplitudes and a better signal-to-noise ratio.

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.

Auditory brain stem responses evoked by lateralized clicks: is lateralization extracted in the human brain stem?

The dependence of binaurally evoked auditory brain stem responses and the binaural difference potential on simultaneously presented interaural time and level differences is investigated in order to assess the representation of stimulus lateralization in the brain stem. Auditory brain stem responses to binaural click stimuli with all combinations of three interaural time and three interaural level differences were recorded from 12 subjects and 4 channels. The latency of Jewett wave V is shortest for zero interaural time difference and longest for the trading stimuli. The amplitude of wave V is largest for centrally perceived stimuli, i.e., the diotic and trading stimuli, and smallest for the most laterally perceived stimuli. The latency of the most prominent peak of the binaural difference potential DN1 mainly depends on the interaural time difference. The amplitude of the components of the binaural difference potential, DP1−DN1, depends similarly on stimulus conditions as wave V amplitude in the case of the binaural stimuli: smallest amplitudes are found for the most lateral stimuli and largest amplitudes for central stimuli. The results demonstrate that interaural level and time differences are not processed independently. This supports the hypothesis that directional information in humans is already extracted and represented at the level of the brain stem.

Interaural delay-dependent changes in the binaural difference potential in cat auditory brainstem response: implications about the origin of the binaural interaction component

Auditory brainstem responses (ABRs) evoked by dichotic clicks with 12 different interaural delays (ITDs) between 0 and 1500 μs were recorded from the vertices of 10 cats under ketamine anesthesia. The so-called binaural difference potential (BDP), considered to be an indicator of binaural interaction (BI), was computed by subtracting the sum of the two monaural responses from the binaural one. The earliest and most prominent component of BDP was a negative deflection (DN1) at a latency between 4 and 4.8 ms. Like all the other components of BDP, DN1 was also due to binaural reduction rather than enhancement of the corresponding ABR wave, P4 in this case. Furthermore, the way its latency increased as a function of ITD was also not compatible with what would be predicted by the delay-line coincidence detector models based on the excitatory-excitatory units in the medial superior olive (MSO). We therefore proposed an alternative hypothesis for the origin of this BI component based on the inhibitory-excitatory (IE) units in the lateral superior olive (LSO). The computational model designed closely simulated the ITD-dependent attenuation and latency shifts observed in DN1. It was therefore concluded that the origin of this BI component in the cat's vertex-ABR could be the lateral lemniscal output of the LSO, although the delay lines which have been shown to exist also in the mammalian brain may play an important role in encoding ITDs. ©1997 Elsevier Science B.V. All rights reserved.

Cellular generators of the binaural difference potential in cat

In humans, lateralization and fusion of binaurally presented clicks are correlated with the latency and amplitude of the binaural difference potential (BDP) (e.g., Furst et al., 1985). The BDP is derived by subtracting the brainstem auditory evoked potential (BAEP) for binaural stimulation from the sum of the BAEPs for left and right monaural stimulation. Our aim in this work was to determine the cellular generators of the BDP and thus identify cells that may be crucial for specific types of binaural sound processing. To this end, we injected kainic acid into the superior olivary complex (SOC) or the cochlear nucleus (CN) in cats and examined the effects of the resulting lesions on the click-evoked BDP. Lesions confined to the anterior anteroventral CN (AVCNa) substantially reduced the BDP, while lesions primarily involving more posterior parts of the CN had little or no effect. BDP reductions occurred for lesions involving either high (> 10 kHz) or lower (< 10 kHz) characteristic frequency (CF) regions of the AVCNa (as well as the posterior CN). Lesions involving the SOC reduced the BDP and, in one case, eliminated the high-pass filtered (270 Hz cutoff) BDP. Combining these results with published information about the physiology and anatomy of auditory brainstem cells, we conclude that: (1) spherical cells in the AVCNa are essential for BDP production, (2) the earliest part of the BDP is generated by medial superior olive (MSO) principal cells which receive spherical cell inputs, (3) a later part is probably generated by the cellular targets of MSO principal cells and, (4) the cells involved in BDP generation have CFs above, as well as below, 10 kHz. Since humans, like cats, have a well-developed MSO, we suggest that the MSO may also be essential for BDP production in humans. Thus, perceptual correlates of the BDP, binaural fusion and click lateralization, apparently involve the MSO.

Electrophysiological correlates of the lateralization of a sound and its echo in human

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.

Field potentials in rat brain stem during binaural acoustic stimulation

Electric field potentials were measured in rat brain stem while presenting monaural or binaural acoustic click stimuli. The binaural difference (BD) potential, defined as the sum of field potentials due to menaural stimuli minus that due to a binaural stimulus, was recorded at 1-mm intervals along a track passing through the inferior colliculus and the superior olivary nuclei. Scalp surface potentials were recorded simultaneously. Prominent BD potentials were observed at all intracranial locations. Scalp surface BD was contemporaneous with an intracranial BD waveform that reversed sign at a level midway between the inferior colliculus and the olivary nuclei. The onset of BD potentials at or below this level occurs 1 ms after the appearance of prominent monaural and binaural response waveforms. This latency difference suggests the existence of an additional synapse in the pathway leading to the structure responsible for BD generation compared to pathways leading to other structures in the superior olivary complex. [Work supported by Penn Lions Hearing Research Foundation.]

Binaural interaction in brainstem auditory evoked potentials elicited by frequency-specific stimuli.

The frequency specificity of the binaural interaction in brainstem auditory evoked potentials (BAEP) was investigated in ten normal-hearing young adults. A novel stimulus paradigm was devised to reduce the influence of the acoustic reflex (middle ear muscle contraction) on the BAEP, and to minimize the effect of variations in noise level. Sequences of six stimuli (rarefaction clicks or Gaussian-shaped tone pulses with carrier frequencies of 1, 2, 4 and 6 kHz) were periodically presented in the following order: right monaural, left monaural, binaural, left monaural, right monaural, binaural, with an interstimulus interval of 22 ms. Since the sequence of monaural stimuli with binaural stimuli interposed produces a uniform loudness and since the acoustic reflex is a consensual reflex, the relative high stimulus repetition rate (approx. 45/s) causes a muscle contraction which is equal on both sides and rather constant in time. This paradigm turned out to be usable for stimulus intensities as high as 80 dB nHL. The binaural difference potential (BDP) was computed by subtracting the sum of the monaurally (ipsilateral and contralateral) evoked potentials from the binaurally evoked potential. The major binaural interaction occurred in the latency range of BAEP waves V and VI, and there was no evidence of interaction in the earlier portion of the BAEP. Both latency and amplitude of the BDP components were evaluated statistically. The latency of the BDP components - except of the lasted one - showed an almost linear dependence both on stimulus intensity and stimulus frequency. The amplitude grew larger with decreasing frequency, and the visual detection threshold elevated as the stimulus frequency increased. Click stimuli, however, produced the largest amplitudes with lowest visual detection threshold. This novel stimulus paradigm appears to be most suitable for routine clinical investigations since high stimulus intensities can be used.

Intra-and extracranially recorded auditory evoked potentials in the cat. I. Source location and binaural interaction

Auditory field potentials (AFPs) were recorded stereotactically from the superior olivary complex and from the exposed inferior colliculus in the Nembutal anaesthetized cat. The brain-stem evoked response (BSER) was recorded simultaneously between an electrode on the dura mater at the vertex and an electrode on one bulla or in the neck musculature. A closed condenser microphone sound system was used to deliver monaural and binaural clicks. The binaural difference potential (BDP) was calculated by subtracting the sum of both monaurally evoked potentials from the binaurally evoked potential. The first binaural interaction was a reduction of BSER wave P4, arising in the superior olivary complex. The large extracellular AFPs generated in both inferior colliculus and superior olivary complex do not correspond to BSER waves, whereas the small volleys preceding the main waves do. These small waves show a much smaller change with recording distance than do the main AFP waves, i.e., the sources and sinks generating these waves appear to be more widely separated. We suggest that, in contrast to extracellular field potentials, the BSER is generated by action potentials in fibre tracts rather than postsynaptic potentials in nuclei. The implications for stimulation and recording laterality of late BSER waves are discussed.