Auditory Brainstem Models: Adapting Cochlear Nuclei Improve Spatial Encoding by the Medial Superior Olive in Reverberation

Listeners typically perceive a sound as originating from the direction of its source, even as direct sound is followed milliseconds later by reflected sound from multiple different directions. Early-arriving sound is emphasised in the ascending auditory pathway, including the medial superior olive (MSO) where binaural neurons encode the interaural-time-difference (ITD) cue for spatial location. Perceptually, weighting of ITD conveyed during rising sound energy is stronger at 600 Hz than at 200 Hz, consistent with the minimum stimulus rate for binaural adaptation, and with the longer reverberation times at 600 Hz, compared with 200 Hz, in many natural outdoor environments. Here, we computationally explore the combined efficacy of adaptation prior to the binaural encoding of ITD cues, and excitatory binaural coincidence detection within MSO neurons, in emphasising ITDs conveyed in early-arriving sound. With excitatory inputs from adapting, nonlinear model spherical bushy cells (SBCs) of the bilateral cochlear nuclei, a nonlinear model MSO neuron with low-threshold potassium channels reproduces the rate-dependent emphasis of rising vs. peak sound energy in ITD encoding; adaptation is equally effective in the model MSO. Maintaining adaptation in model SBCs, and adjusting membrane speed in model MSO neurons, ‘left’ and ‘right’ populations of computationally efficient, linear model SBCs and MSO neurons reproduce this stronger weighting of ITD conveyed during rising sound energy at 600 Hz compared to 200 Hz. This hemispheric population model demonstrates a link between strong weighting of spatial information during rising sound energy, and correct unambiguous lateralisation of a speech source in reverberation.     

Models of Brainstem Responses to Bilateral Electrical Stimulation

A simple, biophysically specified cell model is used to predict responses of binaurally sensitive neurons to patterns of input spikes that represent stimulation by acoustic and electric waveforms. Specifically, the effects of changes in parameters of input spike trains on model responses to interaural time difference (ITD) were studied for low-frequency periodic stimuli, with or without amplitude modulation. Simulations were limited to purely excitatory, bilaterally driven cell models with basic ionic currents and multiple input fibers. Parameters explored include average firing rate, synchrony index, modulation frequency, and latency dispersion of the input trains as well as the excitatory conductance and time constant of individual synapses in the cell model. Results are compared to physiological recordings from the inferior colliculus (IC) and discussed in terms of ITD-discrimination abilities of listeners with cochlear implants. Several empirically observed aspects of ITD sensitivity were simulated without evoking complex neural processing. Specifically, our results show saturation effects in rate–ITD curves, the absence of sustained responses to high-rate unmodulated pulse trains, the renewal of sensitivity to ITD in high-rate trains when inputs are amplitude-modulated, and interactions between envelope and fine-structure delays for some modulation frequencies.     

Sensitivity for interaural time and intensity difference of auditory midbrain neurons in the grassfrog

The sensitivity for interaural time (ITD) and intensity (IID) difference was investigated for single units in the auditory midbrain of the grassfrog. A temporally structured stimulus was used which was presented by means of a closed sound system. At best frequency (BF) the majority of units was selective for ITD as indicated by an asymmetrically (73%) or symmetrically (7%) shaped ITD-rate histogram. About 20% appeared to be nonselective. Units with a symmetrical rate histogram had BFs well above 0.9 kHz, whereas for the other categories no relationship with BF was observed. Most units had a selectivity for ITD which was rather independent from frequency and absolute intensity level. In 62% of the units interaural time difference could be traded by interaural intensity difference. In most cases this so-called time-intensity trading could be explained by the intensity-latency characteristics of auditory nerve fibres. About 20% was sensitive to IID only and 5% to ITD only. A binaural model is proposed which is based on the intensity-rate and intensity-latency characteristics of auditory nerve fibres, the linear summation of excitatory and inhibitory post synaptic potentials in second order neurons, and spatiotemporal integration at the level of third order neurons. By variation of only a small number of parameters, namely strengths and time constants of the connectivities, the range of experimentally observed response patterns could be reproduced.     

Responses of Auditory Nerve and Anteroventral Cochlear Nucleus Fibers to Broadband and Narrowband Noise: Implications for the Sensitivity to Interaural Delays

The quality of temporal coding of sound waveforms in the monaural afferents that converge on binaural neurons in the brainstem limits the sensitivity to temporal differences at the two ears. The anteroventral cochlear nucleus (AVCN) houses the cells that project to the binaural nuclei, which are known to have enhanced temporal coding of low-frequency sounds relative to auditory nerve (AN) fibers. We applied a coincidence analysis within the framework of detection theory to investigate the extent to which AVCN processing affects interaural time delay (ITD) sensitivity. Using monaural spike trains to a 1-s broadband or narrowband noise token, we emulated the binaural task of ITD discrimination and calculated just noticeable differences (jnds). The ITD jnds derived from AVCN neurons were lower than those derived from AN fibers, showing that the enhanced temporal coding in the AVCN improves binaural sensitivity to ITDs. AVCN processing also increased the dynamic range of ITD sensitivity and changed the shape of the frequency dependence of ITD sensitivity. Bandwidth dependence of ITD jnds from AN as well as AVCN fibers agreed with psychophysical data. These findings demonstrate that monaural preprocessing in the AVCN improves the temporal code in a way that is beneficial for binaural processing and may be crucial in achieving the exquisite sensitivity to ITDs observed in binaural pathways.     

Interaural Time Difference Perception with a Cochlear Implant and a Normal Ear

Currently there is a growing population of cochlear-implant (CI) users with (near) normal hearing in the non-implanted ear. This configuration is often called SSD (single-sided deafness) CI. The goal of the CI is often to improve spatial perception, so the question raises to what extent SSD CI listeners are sensitive to interaural time differences (ITDs). In a controlled lab setup, sensitivity to ITDs was investigated in 11 SSD CI listeners. The stimuli were 100-pps pulse trains on the CI side and band-limited click trains on the acoustic side. After determining level balance and the delay needed to achieve synchronous stimulation of the two ears, the just noticeable difference in ITD was measured using an adaptive procedure. Seven out of 11 listeners were sensitive to ITDs, with a median just noticeable difference of 438 μs. Out of the four listeners who were not sensitive to ITD, one listener reported binaural fusion, and three listeners reported no binaural fusion. To enable ITD sensitivity, a frequency-dependent delay of the electrical stimulus was required to synchronize the electric and acoustic signals at the level of the auditory nerve. Using subjective fusion measures and refined by ITD sensitivity, it was possible to match a CI electrode to an acoustic frequency range. This shows the feasibility of these measures for the allocation of acoustic frequency ranges to electrodes when fitting a CI to a subject with (near) normal hearing in the contralateral ear.     

Neuronal sensitivity to interaural time differences in the sound envelope in the auditory cortex of the pallid bat

Interaural time differences in the envelope of a sound (envelope ITDs) can potentially provide spatial information at high frequencies where interaural phase differences (IPDs) are not available. Interaural intensity differences (IIDs) also provide important spatial information at high frequencies. Both IIDs and envelope ITDs can influence spatial perception at high frequencies, but behavioral and physiological studies suggest that IIDs dominate perception. This study examines envelope ITD sensitivity in the auditory cortex of the pallid bat, a species that uses passive sound localization at the low end of its audible range to find prey. Its auditory system is entirely 'high-frequency' in that phase-locking does not occur at the low end of its audible range. If the bat uses ITDs, they must be derived from the envelope of the signal. A previous study of envelope ITD sensitivity in its inferior colliculus (IC) reported that neurons are sensitive to the small +/-70 micros range of available ITDs. This study extends these findings to the cortical level to assess the transformation of ITD sensitivity and the binaural response properties that underlie this sensitivity. Two measures of sensitivity were used. The dynamic ITD range measures the range of ITDs over which the maximum response of a neuron decreases by 80%. When presented with square-wave amplitude-modulated tones statically delayed in arrival time, the average dynamic ITD range in the IC is 304 micros, but dropped to 175 micros in auditory cortex. IC neurons average a 38% change in maximum response over the relevant ITD range, while cortical neurons average a 67% change. Also measured were time-intensity trading ratios, which index the extent to which a change in IID can cause a shift the dynamic ITD range. Average trading ratios are approximately the same in the IC and auditory cortex (17.9 micros/dB vs. 16.7 micros/dB, respectively). Binaural interactions changed from the IC to auditory cortex. In IC, ITD sensitivity is an inhibitory, subtractive process in which ITDs reduce the response evoked by contralateral monaural stimulation. In the auditory cortex, both binaural inhibition and facilitation occur. In the majority of cortical neurons, IID and ITD functions were remarkably similar in shape, having stepped, step-peaked or peaked functions. The binaural interactions (inhibition and/or facilitation) evoked by ITDs and IIDs were also typically similar. These results suggest that IIDs and envelope ITDs are having similar effects on output of the same binaural comparator system.     

Detection of interaural correlation by neurons in the superior olivary complex, inferior colliculus and auditory cortex of the unanesthetized rabbit

A critical binaural cue important for sound localization and detection of signals in noise is the interaural time difference (ITD), or difference in the time of arrival of sounds at each ear. The ITD can be determined by cross-correlating the sounds at the two ears and finding the ITD where the correlation is maximal. The amount of interaural correlation is affected by properties of spaces and can therefore be used to assess spatial attributes. To examine the neural basis for sensitivity to the overall level of the interaural correlation, we identified subcollicular neurons and neurons in the inferior colliculus (IC) and auditory cortex of unanesthetized rabbits that were sensitive to ITDs and examined their responses as the interaural correlation was varied. Neurons at each brain level could show linear or non-linear responses to changes in interaural correlation. The direction of the non-linearities in most neurons was to increase the slope of the response change for correlations near 1.0. The proportion of neurons with non-linear responses was similar in subcollicular and IC neurons but increased in the auditory cortex. Non-linear response functions to interaural correlation were not related to the type of response as determined by the tuning to ITDs across frequencies. The responses to interaural correlation were also not related to the frequency tuning of the neuron, unlike the responses to ITD, which broadens for neurons tuned to lower frequencies. The neural discriminibility of the ITD using frozen noise in the best neurons was similar to the behavioral acuity in humans at a reference correlation of 1.0. However, for other reference ITDs the neural discriminibility was more linear and generally better than the human discriminibility of the interaural correlation, suggesting that stimulus rather than neural variability is the basis for the decline in human performance at lower levels of interaural correlation.     

Sensitivity to Interaural Time Differences Conveyed in the Stimulus Envelope: Estimating Inputs of Binaural Neurons Through the Temporal Analysis of Spike Trains

Sound-source localization in the horizontal plane relies on detecting small differences in the timing and level of the sound at the two ears, including differences in the timing of the modulated envelopes of high-frequency sounds (envelope interaural time differences (ITDs)). We investigated responses of single neurons in the inferior colliculus (IC) to a wide range of envelope ITDs and stimulus envelope shapes. By a novel means of visualizing neural activity relative to different portions of the periodic stimulus envelope at each ear, we demonstrate the role of neuron-specific excitatory and inhibitory inputs in creating ITD sensitivity (or the lack of it) depending on the specific shape of the stimulus envelope. The underlying binaural brain circuitry and synaptic parameters were modeled individually for each neuron to account for neuron-specific activity patterns. The model explains the effects of envelope shapes on sensitivity to envelope ITDs observed in both normal-hearing listeners and in neural data, and has consequences for understanding how ITD information in stimulus envelopes might be maximized in users of bilateral cochlear implants—for whom ITDs conveyed in the stimulus envelope are the only ITD cues available.     

Measuring and modelling the response of auditory midbrain neurons in the grassfrog to temporally structured binaural stimuli

The combined selectivity for amplitude modulation frequency (AMF) and interaural time difference (ITD) was investigated for single units in the auditory midbrain of the grassfrog. Stimuli were presented by means of a closed sound system. A large number of units was found to be selective for AMF (95%) or ITD (85%) and mostly, these selectivities were intricately coupled. At zero ITD most units showed a band-pass (54%) or bimodal (24%) AMF-rate histogram. At an AMF of 36 Hz, which is equal to the pulse repetition rate of the mating call, 70% of the units possessed an asymmetrical ITD-rate histogram, whereas about 15% showed a symmetrically peaked histogram. With binaural stimulation more units appeared to be selective for AMF (95%) as was the case with monaural stimulation (85%). A large fraction of the units appeared to be most selective for ITD at AMFs of 36 and 72 Hz, whereas units seldomly exhibited ITD selectivity with unmodulated tones. Based upon previous papers (Melssen et al., 1990; Van Stokkum, 1990) a binaural model is proposed to explain these findings. An auditory midbrain neuron is modelled as a third order neuron which receives excitatory input from second order neurons. Furthermore the model neuron receives inputs from the other ear, which may be either excitatory or inhibitory. Spatiotemporal integration of inputs from both ears, followed by action potential generation, produces a combined selectivity for AMF and ITD. In particular the responses of an experimentally observed EI neuron to a set of stimuli are reproduced well by the model.     

Tuning to Binaural Cues in Human Auditory Cortex

Interaural level and time differences (ILD and ITD), the primary binaural cues for sound localization in azimuth, are known to modulate the tuned responses of neurons in mammalian auditory cortex (AC). The majority of these neurons respond best to cue values that favor the contralateral ear, such that contralateral bias is evident in the overall population response and thereby expected in population-level functional imaging data. Human neuroimaging studies, however, have not consistently found contralaterally biased binaural response patterns. Here, we used functional magnetic resonance imaging (fMRI) to parametrically measure ILD and ITD tuning in human AC. For ILD, contralateral tuning was observed, using both univariate and multivoxel analyses, in posterior superior temporal gyrus (pSTG) in both hemispheres. Response-ILD functions were U-shaped, revealing responsiveness to both contralateral and—to a lesser degree—ipsilateral ILD values, consistent with rate coding by unequal populations of contralaterally and ipsilaterally tuned neurons. In contrast, for ITD, univariate analyses showed modest contralateral tuning only in left pSTG, characterized by a monotonic response-ITD function. A multivoxel classifier, however, revealed ITD coding in both hemispheres. Although sensitivity to ILD and ITD was distributed in similar AC regions, the differently shaped response functions and different response patterns across hemispheres suggest that basic ILD and ITD processes are not fully integrated in human AC. The results support opponent-channel theories of ILD but not necessarily ITD coding, the latter of which may involve multiple types of representation that differ across hemispheres.     

Interaural Time-Delay Sensitivity in Bilateral Cochlear Implant Users: Effects of Pulse Rate, Modulation Rate, and Place of Stimulation

Electrical interaural time delay (ITD) discrimination was measured using 300-ms bursts applied to binaural pitch matched electrodes at basal, mid, and apical locations in each ear. Six bilateral implant users, who had previously shown good ITD sensitivity at a pulse rate of 100 pulses per second (pps), were assessed. Thresholds were measured as a function of pulse rate between 100 and 1,000 Hz, as well as modulation rate over that same range for high-rate pulse trains at 6,000 pps. Results were similar for all three places of stimulation and showed decreasing ITD sensitivity as either pulse rate or modulation rate increased, although the extent of that effect varied across subjects. The results support a model comprising a common ITD mechanism for high- and low-frequency places of stimulation, which, for electrical stimulation, is rate-limited in the same way across electrodes because peripheral temporal responses are largely place invariant. Overall, ITD sensitivity was somewhat better with unmodulated pulse trains than with high-rate pulse trains modulated at matched rates, although comparisons at individual rates showed that difference to be significant only at 300 Hz. Electrodes presenting with the lowest thresholds at 600 Hz were further assessed using bursts with a ramped onset of 10 ms. The slower rise time resulted in decreased performance in four of the listeners, but not in the two best performers, indicating that those two could use ongoing cues at 600 Hz. Performance at each place was also measured using single-pulse stimuli. Comparison of those data with the unmodulated 300-ms burst thresholds showed that on average, the addition of ongoing cues beyond the onset enhanced overall ITD sensitivity at 100 and 300 Hz, but not at 600 Hz. At 1,000 Hz, the added ongoing cues actually decreased performance. That result is attributed to the introduction of ambiguous cues within the physiologically relevant range and increased dichotic firing.     

Congenital and Prolonged Adult-Onset Deafness Cause Distinct Degradations in Neural ITD Coding with Bilateral Cochlear Implants

Bilateral cochlear implant (CI) users perform poorly on tasks involving interaural time differences (ITD), which are critical for sound localization and speech reception in noise by normal-hearing listeners. ITD perception with bilateral CI is influenced by age at onset of deafness and duration of deafness. We previously showed that ITD coding in the auditory midbrain is degraded in congenitally deaf white cats (DWC) compared to acutely deafened cats (ADC) with normal auditory development (Hancock et al., J. Neurosci, 30:14068). To determine the relative importance of early onset of deafness and prolonged duration of deafness for abnormal ITD coding in DWC, we recorded from single units in the inferior colliculus of cats deafened as adults 6 months prior to experimentation (long-term deafened cats, LTDC) and compared neural ITD coding between the three deafness models. The incidence of ITD-sensitive neurons was similar in both groups with normal auditory development (LTDC and ADC), but significantly diminished in DWC. In contrast, both groups that experienced prolonged deafness (LTDC and DWC) had broad distributions of best ITDs around the midline, unlike the more focused distributions biased toward contralateral-leading ITDs present in both ADC and normal-hearing animals. The lack of contralateral bias in LTDC and DWC results in reduced sensitivity to changes in ITD within the natural range. The finding that early onset of deafness more severely degrades neural ITD coding than prolonged duration of deafness argues for the importance of fitting deaf children with sound processors that provide reliable ITD cues at an early age.     

Learning to discriminate interaural time differences: an exploratory study with amplitude-modulated stimuli

The advent of bilateral cochlear implants (CIs) has increased interest in learning on binaural tasks, and studies in normal-hearing listeners provide important background information. However, few studies have considered learning with discrimination of interaural time difference (ITD). Here, learning with ITD was explored using stimuli that are more relevant to bilateral CIs than used previously. Inexperienced listeners were trained with envelope-based ITD using high-frequency amplitude-modulated tones with or without an interaural carrier frequency difference (IFD), the former to simulate asymmetrical bilateral CI insertions. All were tested with and without IFD before and after training. In most listeners, ITD thresholds improved substantially with training, not necessarily reaching asymptote after 3,000 trials. In these, the magnitude and time-course of learning was larger than anticipated from a previous study with low-frequency ITD. Learning generalized across IFD and the effect of IFD on ITD thresholds at post-test was smaller than reported previously. These results have implications for studies of bilateral CIs, such as the need to provide extensive training to avoid over-estimating any apparent 'impairment'.     

Improved Neural Coding of ITD with Bilateral Cochlear Implants by Introducing Short Inter-pulse Intervals

Bilateral cochlear implant (CI) users have poor perceptual sensitivity to interaural time differences (ITDs), which limits their ability to localize sounds and understand speech in noisy environments. This is especially true for high-rate (>?300 pps) periodic pulse trains, which are used as carriers in CI processors. Here, we investigate a novel stimulation strategy in which extra pulses are added to high-rate periodic pulse trains to introduce short inter-pulse intervals (SIPIs). We hypothesized that SIPIs can improve neural ITD sensitivity similarly to the effect observed by randomly jittering IPIs (Hancock et al., J. Neurophysiol. 108:714–28, 2012). To test this hypothesis, we measured ITD sensitivity of single units in the inferior colliculus (IC) of unanesthetized rabbits with bilateral CIs. Introducing SIPIs into high-rate pulse trains significantly increased firing rates for ~?60 % of IC neurons, and the extra spikes tended to be synchronized to the SIPIs. The additional firings produced by SIPIs uncovered latent ITD sensitivity that was comparable to that observed with low-rate pulse trains. In some neurons, high spontaneous firing rates masked the ITD sensitivity introduced by SIPIs. ITD sensitivity in these neurons could be revealed by emphasizing stimulus-synchronized spikes with a coincidence detection analysis. Overall, these results with SIPIs are consistent with the effects observed previously with jittered pulse trains, with the added benefit of retaining control over the timing and number of SIPIs. A novel CI processing strategy could incorporate SIPIs by inserting them at selected times to high-rate pulse train carriers. Such a strategy could potentially improve ITD perception without degrading speech intelligibility and thereby improve outcomes for bilateral CI users.     

Processing Temporal Modulations in Binaural and Monaural Auditory Stimuli by Neurons in the Inferior Colliculus and Auditory Cortex

Processing dynamic changes in the stimulus stream is a major task for sensory systems. In the auditory system, an increase in the temporal integration window between the inferior colliculus (IC) and auditory cortex is well known for monaural signals such as amplitude modulation, but a similar increase with binaural signals has not been demonstrated. To examine the limits of binaural temporal processing at these brain levels, we used the binaural beat stimulus, which causes a fluctuating interaural phase difference, while recording from neurons in the unanesthetized rabbit. We found that the cutoff frequency for neural synchronization to the binaural beat frequency (BBF) decreased between the IC and auditory cortex, and that this decrease was associated with an increase in the group delay. These features indicate that there is an increased temporal integration window in the cortex compared to the IC, complementing that seen with monaural signals. Comparable measurements of responses to amplitude modulation showed that the monaural and binaural temporal integration windows at the cortical level were quantitatively as well as qualitatively similar, suggesting that intrinsic membrane properties and afferent synapses to the cortical neurons govern the dynamic processing. The upper limits of synchronization to the BBF and the band-pass tuning characteristics of cortical neurons are a close match to human psychophysics.     

小鼠中脑下丘神经元对单、双耳听觉信息突触反应的多样性研究

目的研究小鼠中脑下丘单个神经元对单耳对侧、同侧声刺激的突触反应及双侧同时刺激的整合反应,探索其潜在的神经生理学基础与神经环路。方法52只正常C57小鼠,采用声刺激系统记录下丘单个神经元对单耳对侧、单耳同侧声刺激的频率-幅度反应域(frequency-amplitude response areas,FARA),获得神经元的特征频率(characteristic frequency,CF)和最低阈值(minimum threshold,MT)。采用活体全细胞膜片钳技术,在最适声刺激条件下(即声音参数为CF与MT),记录下丘同一个神经元分别对单耳对侧、单耳同侧声刺激的突触反应,以及双侧声音同时刺激时的突触整合反应,对记录到的突触反应及整合反应进行分类分析。结果共记录到146个下丘神经元,CF-对侧与CF-同侧分别为14.9±4.8、14.7±5.0 kHz,两者成直线相关且相关系数为1.0258;MT-对侧为19.3±19.3 dB,显著低于MT-同侧(45.1±18.6 dB),差异有统计学意义(P<0.001)。根据神经元对单耳刺激的反应是兴奋(excitation,E)、无反应(no response,O)还是抑制(inhibition,I),可将这146个神经元分成7种不同类型,即EE、EO、EI、II、IO、IE和CM(complex-mode,CM)双耳神经元,分别占66.4%(97/146)、15.8%(23/146)、4.1%(6/146)、6.8%(10/146)、1.4%(2/146)、1.4%(2/146)和4.1%(6/146)。根据双耳整合特性,EE神经元可分成抑制EE/I、易化EE/F和无整合EE/N三种类型,分别占38.1%(37/97)、20.6%(20/97)和41.2%(40/97)。EO和II神经元对双耳信息的整合反应可出现抑制和无整合两种,EI神经元仅表现出抑制的整合反应,而IO和IE神经元则呈双耳无整合效应。结论EE、EO、EI、II、IO、IE和CM这7种不同类型的下丘双耳神经元各自具有不同的双侧突触结构和神经环路。     

Coding of interaural time differences of transients in auditory cortex of Rattus norvegicus: implications for the evolution of mammalian sound localization.

We obtained quantitative evidence on the coding of interaural time differences (ITDs) of click stimuli by 40 single neurons in the auditory cortex of anesthetized albino rats. Most of the neurons (31/40) received an excitatory input from the contralateral ear, and an inhibitory input from the ipsilateral ear (EI cells). These neurons expressed their sensitivity to ITDs in a sigmoidal relation between spike count and ITD, with maximal responses associated with contralateral-leading ITDs. The mean ITD dynamic range was 590 microseconds. The dynamic ranges typically encompassed at least part of the behaviorally-relevant range (about +/- 130 microseconds). Variations in ITD from 130 microseconds favoring one ear to 130 microseconds favoring the other ear caused spike response rate changes, on average, of 29.5%. These data are similar to those previously presented for the central auditory systems of larger mammals, whose auditory localization acuity is significantly better than that of the rat. We argue, therefore, that the sound localization mechanisms based on transient ITDs have not evolved in a fashion that covaries with interaural distance, and that there exists a mismatch between the ITDs the rat will encounter in the free field, and the ITDs which are encoded by its nervous system. This may be one reason why sound localization acuity has a roughly inverse relation to interaural distance.     

Spatiotemporal representation of binaural difference in time and intensity of sound in the guinea pig auditory cortex.

Neural activity of the auditory cortex (AC) in response to a change of interaural intensity difference (IID) and interaural time difference (ITD) of sound stimuli was observed by optical recording with a 12 x 12 photodiode array and the voltage-sensitive dye, RH795. Guinea pigs (280-450 g) were anesthetized with sodium pentobarbital (30 mg/kg) and supplemental doses of neuroleptic solutions. When both ears were stimulated dichotically by tone bursts (14 kHz, 75 dB SPL), excitatory optical signals appeared in both anterior (A) and dorsocaudal (DC) fields of AC. An increase of intensity of ipsilateral stimulation from 65 to 95 dB SPL caused a decrease of neural activity of isofrequency bands in both fields. An increase of ipsilateral leads from -2.5 to 10 ms resulted in a gradual decrease of the amplitude of the excitatory responses. A strong inhibition was observed in field DC and the ventral portion of field A. These results show the different spatiotemporal representation of IID and ITD sensitivities in AC. However, the ipsilateral lead inducing a large inhibition was much longer than the time difference (80 micros) calculated from the interaural distance of the guinea pig. This indicates that the longer binaural inhibition observed in AC would have a different functional significance from that of the neural system of ITD detection in the guinea pig.     

The Effect of Interaural Fluctuation Rate on Correlation Change Discrimination

While bilateral cochlear implants (CIs) provide some binaural benefits, these benefits are limited compared to those observed in normal-hearing (NH) listeners. The large frequency-to-electrode allocation bandwidths (BWs) in CIs compared to auditory filter BWs in NH listeners increases the interaural fluctuation rate available for binaural unmasking, which may limit binaural benefits. The purpose of this work was to investigate the effect of interaural fluctuation rate on correlation change discrimination and binaural masking-level differences in NH listeners presented a CI simulation using a pulsed-sine vocoder. In experiment 1, correlation-change just-noticeable differences (JNDs) and tone-in-noise thresholds were measured for narrowband noises with different BWs and center frequencies (CFs). The results suggest that the BW, CF, and/or interaural fluctuation rate are important factors for correlation change discrimination. In experiment 2, the interaural fluctuation rate was systematically varied and dissociated from changes in BW and CF by using a pulsed-sine vocoder. Results indicated that the interaural fluctuation rate did not affect correlation change JNDs for correlated reference noises; however, slow interaural fluctuations increased correlation change JNDs for uncorrelated reference noises. In experiment 3, the BW, CF, and vocoder pulse rate were varied while interaural fluctuation rate was held constant. JNDs increased for increasing BW and decreased for increasing CF. In summary, relatively fast interaural fluctuation rates are not detrimental for detecting changes in interaural correlation. Thus, limiting factors to binaural benefits in CI listeners could be a result of other temporal and/or spectral deficiencies from electrical stimulation.     

The Interaural Time Difference Pathway: a Comparison of Spectral Bandwidth and Correlation Sensitivity at Three Anatomical Levels

Temporal differences between the two ears are critical for spatial hearing. They can be described along axes of interaural time difference (ITD) and interaural correlation, and their processing starts in the brainstem with the convergence of monaural pathways which are tuned in frequency and which carry temporal information. In previous studies, we examined the bandwidth (BW) of frequency tuning at two stages: the auditory nerve (AN) and inferior colliculus (IC), and showed that BW depends on characteristic frequency (CF) but that there is no difference in the mean BW of these two structures when measured in a binaural, temporal framework. This suggested that there is little frequency convergence in the ITD pathway between AN and IC and that frequency selectivity determined by the cochlear filter is preserved up to the IC. Unexpectedly, we found that AN and IC neurons can be similar in CF and BW, yet responses to changes in interaural correlation in the IC were different than expected from coincidence patterns (“pseudo-binaural” responses) in the AN. To better understand this, we here examine the responses of bushy cells, which provide monaural inputs to binaural neurons. Using broadband noise, we measured BW and correlation sensitivity in the cat trapezoid body (TB), which contains the axons of bushy cells. This allowed us to compare these two metrics at three stages in the ITD pathway. We found that BWs in the TB are similar to those in the AN and IC. However, TB neurons were found to be more sensitive to changes in stimulus correlation than AN or IC neurons. This is consistent with findings that show that TB fibers are more temporally precise than AN fibers, but is surprising because it suggests that the temporal information available monaurally is not fully exploited binaurally.