Interaural Time Differences Research Articles

Morphology and unbiased stereology of the lateral superior olive in the short-beaked common dolphin, Delphinus delphis (Cetacea, Delphinidae).

In all mammals, the superior olivary complex (SOC) comprises a group of auditory brainstem nuclei that are important for sound localization. Its principal nuclei, the lateral superior olive (LSO) and the medial superior olive (MSO) process interaural time and intensity differences, which are the main cues for sound localization in the horizontal plane. Toothed whales (odontocetes) rely heavily on hearing and echolocation for foraging, orientation, and communication and localize sound with great acuity. The investigation of the SOC in odontocetes provides insight into adaptations to underwater hearing and echolocation. However, quantitative anatomical data for odontocetes are currently lacking. We quantified the volume, total neuron number, and neuron density of the LSO of six common dolphins (Delphinus delphis) using the Cavalieri principle and the unbiased stereology optical fractionator. Our results show that the LSO in D. delphis has a volume of 150 + (SD=27) mm3 , which is on average 69 (SEM=19) times larger than the LSO in human, or 37 (SEM=11) times larger than the human LSO and MSO combined. The LSO of D. delphis contains 20,876 ± (SD=3300) neurons. In comparison, data reported for the human brainstem indicate the LSO has only about ¼ that number but about the same number for the LSO and MSO combined (21,100). LSO neurons range from 21 to 25 μm (minor axis) and from 44 to 61 μm (major axis) in transverse sections. The LSO neuron packing density is 1080 ± (SD=204) neurons/mm3 , roughly half of the LSO neuron density in human. SMI-32-immunohistochemistry was used to visualize projection neurons in the LSO and revealed the presence of principal, marginal, and multipolar neurons in transverse sections. The distinct morphology of the LSO likely reflects the common dolphin's superb sensitivity to ultra-high frequencies and ability to detect and analyze sounds and their location as part of its underwater spatial localization and echolocation tasks.

The effect of target and interferer frequency on across-frequency binaural interference of interaural-level-difference sensitivity.

Across-frequency binaural interference occurs when the sensitivity to changes in interaural differences in a target sound is decreased by a spectrally remote diotic interfering sound. For interaural time differences (ITDs), low-frequency (e.g., 0.5 kHz) interferers cause more interference on high-frequency (e.g., 4 kHz) targets than vice versa. For interaural level differences (ILDs), however, it is unclear if a frequency dependence exists. Therefore, ILD discrimination thresholds and across-frequency binaural interference were measured for target and interferer frequencies between 0.5 and 8 kHz (for tones) or 12 kHz (for narrowband noises). For tones, 8-kHz targets experienced the least interference and 8-kHz interferers produced the most interference, suggesting that higher-frequency ILDs are a more heavily weighted localization cue than lower-frequency ILDs. For narrowband noises, the frequency-dependent interference patterns increased in complexity in comparison to tones. Low-frequency ITD dominance (from randomly varying onset ITDs) and grouping cues (e.g., envelope modulations) might explain some of the complexity in the interference patterns for the noises. These data contribute to a better understanding of across-frequency ILD processing, which remains poorly understood.

Sound localisation of low- and high-frequency sounds in cochlear implant users with single-sided deafness

Objective Localisation of low- and high-frequency sounds in single-sided deaf cochlear implant users was investigated using noise stimuli designed to mitigate monaural localisation cues. Design Within subject design. Sound source localisation was tested in the horizontal plane using an array of seven loudspeakers along the azimuthal angle span from −90° to +90°. Stimuli were broadband noise and high- and low-frequency noise. Study sample Twelve adult subjects with single-sided deafness participated in the study. All had normal hearing in the healthy ear and were supplied with a cochlear implant (CI) in their deaf ear. Results With broadband noise, the mean angular localisation error was 39° in aided condition as compared to a median angular error of 83.6° when the speech processor was not worn. For high-frequency noise, the median angular error was 30° and for low-frequency noise, it was 46° in the CI-aided condition. Conclusions Single-sided deaf CI users show the best sound localisation for high-frequency sounds. This supports the view that interaural level differences are dominant for sound localisation in these listeners. Nonetheless, a limited ability to localise low-frequency sounds was observed, which may be based on the supportive perception of interaural time differences.

Robust parameter strategy for Wiener-based binaural noise reduction methods in hearing aids

This work presents a method for designing the weighting parameter required for Wiener-based binaural noise reduction methods. This parameter establishes the desired trade-off between noise reduction and binaural cue preservation in hearing aids. The proposed strategy was specially derived for the preservation of the interaural level difference, interaural time difference and interaural coherence binaural cues. It is defined as a function of the average input noise power at the microphones, providing robustness against the influence of joint changes in noise and speech power (Lombard effect), as well as signal-to-noise ratio variations. A theoretical framework based on the mathematical definition of the degree of homogeneity is presented and applied to a generic augmented Wiener-based cost function. The theoretical insights obtained are supported by computational simulations and psychoacoustic experiments using the multichannel Wiener filter with a noise interaural transfer function preservation technique (MWF-ITFv), as a case study. Statistical analysis indicates that, compared to a purely fixed weighting parameter, the proposed structure for the weighting parameter and the design method of its fixed part provide significant robustness against changes in the noise binaural cues at the cost of a small noise reduction performance decrease and speech binaural cue distortion.

Auditory Brain Stem Responses in the C57BL/6J Fragile X Syndrome-Knockout Mouse Model.

Sensory hypersensitivity, especially in the auditory system, is a common symptom in Fragile X syndrome (FXS), the most common monogenic form of intellectual disability. However, linking phenotypes across genetic background strains of mouse models has been a challenge and could underly some of the issues with translatability of drug studies to the human condition. This study is the first to characterize the auditory brain stem response (ABR), a minimally invasive physiological readout of early auditory processing that is also used in humans, in a commonly used mouse background strain model of FXS, C57BL/6J. We measured morphological features of pinna and head and used ABR to measure the hearing range, and monaural and binaural auditory responses in hemizygous males, homozygous females, and heterozygous females compared with those in wild-type mice. Consistent with previous study, we showed no difference in morphological parameters across genotypes or sexes. There was no significant difference in hearing range between the sexes or genotypes, however there was a trend towards high frequency hearing loss in male FXS mice. In contrast, female mice with homozygous FXS had a decreased amplitude of wave IV of the monaural ABR, while there was no difference in males for amplitudes and no change in latency of ABR waveforms across sexes and genotypes. Finally, males with FXS had an increased latency of the binaural interaction component (BIC) at 0 interaural timing difference compared with that in wild-type males. These findings further clarify auditory brain stem processing in FXS by adding more information across genetic background strains allowing for a better understanding of shared phenotypes.

The Relationship Between Interaural Insertion-Depth Differences, Scalar Location, and Interaural Time-Difference Processing in Adult Bilateral Cochlear-Implant Listeners.

Sensitivity to interaural time differences (ITDs) in acoustic hearing involves comparison of interaurally frequency-matched inputs. Bilateral cochlear-implant arrays are, however, only approximately aligned in angular insertion depth and scalar location across the cochleae. Interaural place-of-stimulation mismatch therefore has the potential to impact binaural perception. ITD left-right discrimination thresholds were examined in 23 postlingually-deafened adult bilateral cochlear-implant listeners, using low-rate constant-amplitude pulse trains presented via direct stimulation to single electrodes in each ear. Angular insertion depth and scalar location measured from computed-tomography (CT) scans were used to quantify interaural mismatch, and their association with binaural performance was assessed. Number-matched electrodes displayed a median interaural insertion-depth mismatch of 18° and generally yielded best or near-best ITD discrimination thresholds. Two listeners whose discrimination thresholds did not show this pattern were confirmed via CT to have atypical array placement. Listeners with more number-matched electrode pairs located in the scala tympani displayed better thresholds than listeners with fewer such pairs. ITD tuning curves as a function of interaural electrode separation were broad; bandwidths at twice the threshold minimum averaged 10.5 electrodes (equivalent to 5.9 mm for a Cochlear-brand pre-curved array). Larger angular insertion-depth differences were associated with wider bandwidths. Wide ITD tuning curve bandwidths appear to be a product of both monopolar stimulation and angular insertion-depth mismatch. Cases of good ITD sensitivity with very wide bandwidths suggest that precise matching of insertion depth is not critical for discrimination thresholds. Further prioritizing scala tympani location at implantation should, however, benefit ITD sensitivity.

Measuring and Modeling Cue Dependent Spatial Release from Masking in the Presence of Typical Delays in the Treatment of Hearing Loss.

In asymmetric treatment of hearing loss, processing latencies of the modalities typically differ. This often alters the reference interaural time difference (ITD) (i.e., the ITD at 0° azimuth) by several milliseconds. Such changes in reference ITD have shown to influence sound source localization in bimodal listeners provided with a hearing aid (HA) in one and a cochlear implant (CI) in the contralateral ear. In this study, the effect of changes in reference ITD on speech understanding, especially spatial release from masking (SRM) in normal-hearing subjects was explored. Speech reception thresholds (SRT) were measured in ten normal-hearing subjects for reference ITDs of 0, 1.75, 3.5, 5.25 and 7 ms with spatially collocated (S0N0) and spatially separated (S0N90) sound sources. Further, the cues for separation of target and masker were manipulated to measure the effect of a reference ITD on unmasking by A) ITDs and interaural level differences (ILDs), B) ITDs only and C) ILDs only. A blind equalization-cancellation (EC) model was applied to simulate all measured conditions. SRM decreased significantly in conditions A) and B) when the reference ITD was increased: In condition A) from 8.8 dB SNR on average at 0 ms reference ITD to 4.6 dB at 7 ms, in condition B) from 5.5 dB to 1.1 dB. In condition C) no significant effect was found. These results were accurately predicted by the applied EC-model. The outcomes show that interaural processing latency differences should be considered in asymmetric treatment of hearing loss.

Effects of Hearing Loss on Interaural Time Difference Sensitivity at Low and High Frequencies.

While many studies have reported a loss of sensitivity to interaural time differences (ITDs) carried in the fine structure of low-frequency signals for listeners with hearing loss, relatively few data are available on the perception of ITDs carried in the envelope of high-frequency signals in this population. The relevant studies found stronger effects of hearing loss at high frequencies than at low frequencies in most cases, but small subject numbers and several confounding effects prevented strong conclusions from being drawn. In the present study, we revisited this question while addressing some of the issues identified in previous studies. Participants were ten young adults with normal hearing (NH) and twenty adults with sensorineural hearing impairment (HI) spanning a range of ages. ITD discrimination thresholds were measured for octave-band-wide “rustle” stimuli centered at 500 Hz or 4000 Hz, which were presented at 20 or 40 dB sensation level. Broadband rustle stimuli and 500-Hz pure-tone stimuli were also tested. Thresholds were poorer on average for the HI group than the NH group. The ITD deficit, relative to the NH group, was similar at low and high frequencies for most HI participants. For a small number of participants, however, the deficit was strongly frequency-dependent. These results provide new insights into the binaural perception of complex sounds and may inform binaural models that incorporate effects of hearing loss.

Binaural-cue Weighting and Training-Induced Reweighting Across Frequencies.

During sound lateralization, the information provided by interaural differences in time (ITD) and level (ILD) is weighted, with ITDs and ILDs dominating for low and high frequencies, respectively. For mid frequencies, the weighting between these binaural cues can be changed via training. The present study investigated whether binaural-cue weights change gradually with increasing frequency region, whether they can be changed in various frequency regions, and whether such binaural-cue reweighting generalizes to untrained frequencies. In two experiments, a total of 39 participants lateralized 500-ms, 1/3-octave-wide noise bursts containing various ITD/ILD combinations in a virtual audio-visual environment. Binaural-cue weights were measured before and after a 2-session training in which, depending on the group, either ITDs or ILDs were visually reinforced. In experiment 1, four frequency bands (centered at 1000, 1587, 2520, and 4000 Hz) and a multiband stimulus comprising all four bands were presented during weight measurements. During training, only the 1000-, 2520-, and 4000-Hz bands were presented. In experiment 2, the weight measurements only included the two mid-frequency bands, while the training only included the 1587-Hz band. ILD weights increased gradually from low- to high-frequency bands. When ILDs were reinforced during training, they increased for the 4000- (experiment 1) and 2520-Hz band (experiment 2). When ITDs were reinforced, ITD weights increased only for the 1587-Hz band (at specific azimuths). This suggests that ILD reweighting requires high, and ITD reweighting requires low frequencies without including frequency regions providing fine-structure ITD cues. The changes in binaural-cue weights were independent of the trained bands, suggesting some generalization of binaural-cue reweighting.

Hybrid multi-harmonic model for the prediction of interaural time differences in individual behind-the-ear hearing-aid-related transfer functions

Spatial sound perception in aided listeners partly relies on hearing-aid-related transfer functions (HARTFs), describing the directional acoustic paths between a sound source and the hearing-aid (HA) microphones. Compared to head-related transfer functions (HRTFs), the HARTFs of behind-the-ear HAs exhibit substantial differences in spectro-temporal characteristics and binaural cues such as interaural time differences (ITDs). Since assumptions on antipodal microphone placement on the equator of a three-concentric sphere are violated in such datasets, predicting the ITDs via Kuhn’s simple analytic harmonic model entails excessive errors. Although angular ear-canal offsets have been addressed in an extended Woodworth model, the prediction errors remain large if the frequency range does not comply with the model specifications. Tuned to the previously inaccurately modelled frequency range between 500 Hz and 1.5 kHz, we propose a hybrid multi-harmonic model to predict the ITDs in HRTFs and HARTFs for arbitrary directions in the horizontal plane with superior accuracy. The target model coefficients are derived from individual directional measurements of 30 adults, wearing two dual-microphone behind-the-ear HAs and two in-ear microphones. Model individualisation is facilitated by the availability of polynomial weights that are applied to subsets of individual anthropometric and HA features to estimate the target model coefficients. The model is published as part of the Auditory Modeling Toolbox (AMT,pausch2022) and supplemented with the individual features and directional datasets.

Deep neural network models of sound localization reveal how perception is adapted to real-world environments

Mammals localize sounds using information from their two ears. Localization in real-world conditions is challenging, as echoes provide erroneous information, and noises mask parts of target sounds. To better understand real-world localization we equipped a deep neural network with human ears and trained it to localize sounds in a virtual environment. The resulting model localized accurately in realistic conditions with noise and reverberation. In simulated experiments, the model exhibited many features of human spatial hearing: sensitivity to monaural spectral cues and interaural time and level differences, integration across frequency, biases for sound onsets, and limits on localization of concurrent sources. But when trained in unnatural environments without either reverberation, noise, or natural sounds, these performance characteristics deviated from those of humans. The results show how biological hearing is adapted to the challenges of real-world environments and illustrate how artificial neural networks can reveal the real-world constraints that shape perception.

Interaural-time-difference thresholds for broad band-limited pulses are affected by relative bandwidth not temporal envelope sharpness.

Humans are sensitive to interaural time differences (ITDs) conveyed by slow modulations on high-frequency carrier signals. Sensitivity appears to be affected by temporal envelope sharpness, but it is unclear if there is a limit to which sharpness affects sensitivity. Pulse trains were varied in relative bandwidth (re: critical bandwidths) and center frequency. ITD sensitivity increased with increasing bandwidth. There was no effect of center frequency when relative bandwidths were analyzed, suggesting that the temporal envelope sharpness (concomitantly absolute bandwidth in Hz) did not affect performance. Rather, sensitivity was most easily explained by recruitment of additional auditory channels.

Effects of place of stimulation on the interaural time difference sensitivity in bilateral electrical intracochlear stimulations: Neurophysiological study in a rat model.

We examined the sensitivity of the neurons in the inferior colliculus (IC) in male and female rats to the interaural time differences (ITDs) conveyed in electrical pulse trains. Using bipolar pairs of electrodes that selectively activate the auditory nerve fibers at different intracochlear locations, we assessed whether the responses to electrical stimulation with ITDs in different frequency regions were processed differently. Most well-isolated single units responded to the electrical stimulation in only one of the apical or basal cochlear regions, and they were classified as either apical or basal units. Regardless of the cochlear stimulating location, more than 70% of both apical and basal units were sensitive to ITDs of electrical stimulation. However, the pulse rate dependence of neural ITD sensitivity differed significantly depending on the location of the stimulation. Moreover, ITD discrimination thresholds and the relative incidence of ITD tuning type markedly differed between units activated by apical and basal stimulations. With apical stimulation, IC neurons had a higher incidence of peak-type ITD function, which mostly exhibited the steepest position of the tuning curve within the rat's physiological ITD range of ±160μs and, accordingly, had better ITD discrimination thresholds than those with basal stimulation. These results support the idea that ITD processing in the IC might be determined by functionally segregated frequency-specific pathways from the cochlea to the auditory midbrain.

Dendritic synapse geometry optimizes binaural computation in a sound localization circuit.

Clustering of synapses allows neurons to overcome attenuation of electrical signals at dendrites. However, we show in avian binaural coincidence detectors computing interaural time difference for sound localization that clustering of synapses rather promotes the dendritic attenuation but augments the intensity tolerance of the binaural computations. Using glutamate uncaging, we found in the neurons that synapses were clustered at distal dendritic branches. Modeling revealed that this strengthened sublinear integration within a dendritic tree but enabled the integration of signals from different trees when inputs grow stronger, preventing monoaural output and maintaining the dynamic range of binaural computation. The extent of this clustering differed according to dendritic length and frequency tuning of neurons, being most prominent for long dendrites and low-frequency tuning. This ensures binaural spatial hearing for wide intensity and frequency ranges, highlighting the importance of coupling of synapse geometry with dendritic morphology and input frequency in sensory signal processing.

Reweighting of Binaural Localization Cues in Bilateral Cochlear-Implant Listeners

Normal-hearing (NH) listeners rely on two binaural cues, the interaural time (ITD) and level difference (ILD), for azimuthal sound localization. Cochlear-implant (CI) listeners, however, rely almost entirely on ILDs. One reason is that present-day clinical CI stimulation strategies do not convey salient ITD cues. But even when presenting ITDs under optimal conditions using a research interface, ITD sensitivity is lower in CI compared to NH listeners. Since it has recently been shown that NH listeners change their ITD/ILD weighting when only one of the cues is consistent with visual information, such reweighting might add to CI listeners’ low perceptual contribution of ITDs, given their daily exposure to reliable ILDs but unreliable ITDs. Six bilateral CI listeners completed a multi-day lateralization training visually reinforcing ITDs, flanked by a pre- and post-measurement of ITD/ILD weights without visual reinforcement. Using direct electric stimulation, we presented 100- and 300-pps pulse trains at a single interaurally place-matched electrode pair, conveying ITDs and ILDs in various spatially consistent and inconsistent combinations. The listeners’ task was to lateralize the stimuli in a virtual environment. Additionally, ITD and ILD thresholds were measured before and after training. For 100-pps stimuli, the lateralization training increased the contribution of ITDs slightly, but significantly. Thresholds were neither affected by the training nor correlated with weights. For 300-pps stimuli, ITD weights were lower and ITD thresholds larger, but there was no effect of training. On average across test sessions, adding azimuth-dependent ITDs to stimuli containing ILDs increased the extent of lateralization for both 100- and 300-pps stimuli. The results suggest that low-rate ITD cues, robustly encoded with future CI systems, may be better exploitable for sound localization after increasing their perceptual weight via training.

Effect of gap detection threshold and localisation acuity on spatial release from masking in older adults*

Objective The primary objective of this experiment was to measure the temporal and spatial processing capabilities of older individuals and use statistical models to identify the individual contributions of these temporal and spatial processing capabilities to spatial release from masking (SRM). Design Repeated measures Study sample Twenty-five older listeners with varying degrees of hearing loss participated in this experiment. SRM using the coordinate response measure, gap detection thresholds and localisation acuity for 1/3-octave-wide Gaussian noise bands centred at 500 and 4000 Hz were measured for all the listeners. Results Older listeners had better speech recognition thresholds when target and maskers were spatially separated as compared to when they were co-located. In addition, hearing loss and localisation acuity at 500 Hz were significant predictors in a multiple regression model predicting SRM. However, gap detection thresholds did not significantly contribute to the multiple regression model predicting SRM. Conclusion Based on our data, we conclude that SRM at 30° spatial separation between the target and symmetric maskers is driven by the ability of the individuals to use interaural time difference cues.

Diverse processing underlying frequency integration in midbrain neurons of barn owls.

Emergent response properties of sensory neurons depend on circuit connectivity and somatodendritic processing. Neurons of the barn owl’s external nucleus of the inferior colliculus (ICx) display emergence of spatial selectivity. These neurons use interaural time difference (ITD) as a cue for the horizontal direction of sound sources. ITD is detected by upstream brainstem neurons with narrow frequency tuning, resulting in spatially ambiguous responses. This spatial ambiguity is resolved by ICx neurons integrating inputs over frequency, a relevant processing in sound localization across species. Previous models have predicted that ICx neurons function as point neurons that linearly integrate inputs across frequency. However, the complex dendritic trees and spines of ICx neurons raises the question of whether this prediction is accurate. Data from in vivo intracellular recordings of ICx neurons were used to address this question. Results revealed diverse frequency integration properties, where some ICx neurons showed responses consistent with the point neuron hypothesis and others with nonlinear dendritic integration. Modeling showed that varied connectivity patterns and forms of dendritic processing may underlie observed ICx neurons’ frequency integration processing. These results corroborate the ability of neurons with complex dendritic trees to implement diverse linear and nonlinear integration of synaptic inputs, of relevance for adaptive coding and learning, and supporting a fundamental mechanism in sound localization.

Barn Owl's Auditory Space Map Activity Matching Conditions for a Population Vector Readout to Drive Adaptive Sound-Localizing Behavior.

Space-specific neurons in the owl's midbrain form a neural map of auditory space, which supports sound-orienting behavior. Previous work proposed that a population vector (PV) readout of this map, implementing statistical inference, predicts the owl's sound localization behavior. This model also predicts the frontal localization bias normally observed and how sound-localizing behavior changes when the signal-to-noise ratio varies, based on the spread of activity across the map. However, the actual distribution of population activity and whether this pattern is consistent with premises of the PV readout model on a trial-by-trial basis remains unknown. To answer these questions, we investigated whether the population response profile across the midbrain map in the optic tectum of the barn owl matches these predictions using in vivo multielectrode array recordings. We found that response profiles of recorded subpopulations are sufficient for estimating the stimulus interaural time difference using responses from single trials. Furthermore, this decoder matches the expected differences in trial-by-trial variability and frontal bias between stimulus conditions of low and high signal-to-noise ratio. These results support the hypothesis that a PV readout of the midbrain map can mediate statistical inference in sound-localizing behavior of barn owls.SIGNIFICANCE STATEMENT While the tuning of single neurons in the owl's midbrain map of auditory space has been considered predictive of the highly specialized sound-localizing behavior of this species, response properties across the population remain largely unknown. For the first time, this study analyzed the spread of population responses across the map using multielectrode recordings and how it changes with signal-to-noise ratio. The observed responses support the hypothesis concerning the ability of a population vector readout to predict biases in orienting behaviors and mediate uncertainty-dependent behavioral commands. The results are of significance for understanding potential mechanisms for the implementation of optimal behavioral commands across species.

Sensitivity to Envelope Interaural Time Differences: Modeling Auditory Modulation Filtering.

For amplitude-modulated sound, the envelope interaural time difference (ITDENV) is a potential cue for sound-source location. ITDENV is encoded in the lateral superior olive (LSO) of the auditory brainstem, by excitatory-inhibitory (EI) neurons receiving ipsilateral excitation and contralateral inhibition. Between human listeners, sensitivity to ITDENV varies considerably, but ultimately decreases with increasing stimulus carrier frequency, and decreases more strongly with increasing modulation rate. Mechanisms underlying the variation in behavioral sensitivity remain unclear. Here, with increasing carrier frequency (4-10kHz), as we phenomenologically model the associated decrease in ITDENV sensitivity using arbitrarily fewer neurons consistent across populations, we computationally model the variable sensitivity across human listeners and modulation rates (32-800Hz) as the decreasing range of membrane frequency responses in LSO neurons. Transposed tones stimulate a bilateral auditory-periphery model, driving model EI neurons where electrical membrane impedance filters the frequency content of inputs driven by amplitude-modulated sound, evoking modulation filtering. Calculated from Fisher information in spike-rate functions of ITDENV, for model EI neuronal populations distinctly reflecting the LSO range in membrane frequency responses, just-noticeable differences in ITDENV collectively reproduce the largest variation in ITDENV sensitivity across human listeners. These slow to fast model populations each generally match the best human ITDENV sensitivity at a progressively higher modulation rate, by membrane-filtering and spike-generation properties producing realistically less than Poisson variance. Non-resonant model EI neurons are also sensitive to interaural intensity differences. With peripheral filters centered between carrier frequency and modulation sideband, fast resonant model EI neurons extend ITDENV sensitivity above 500-Hz modulation.

Axonal Conduction Delay Shapes the Precision of the Spatial Hearing in A Spiking Neural Network Model of Auditory Brainstem.

One method by which the mammalian sound localization pathway localizes sound sources is by analyzing the microsecond-level difference between the arrival times of a sound at the two ears. However, how the neural circuits in the auditory brainstem precisely integrate signals from the two ears, and what the underlying mechanisms are, remains to be understood. Recent studies have reported that variations of axon myelination in the auditory brainstem produces various axonal conduction velocities and sophisticated temporal dynamics, which have not been well characterized in most existing models of sound localization circuits. Here, we present a spiking neural network model of the auditory brainstem to investigate how axon myelinations affect the precision of sound localization. Sound waves with different interaural time differences (ITDs) are encoded and used as stimuli, and the axon properties in the network are adjusted, and the corresponding axonal conduction delays are computed with a multi-compartment axon model. Through the simulation, the sensitivity of ITD perception varies with the myelin thickness of axons in the contralateral input pathways to the medial superior olive (MSO). The ITD perception becomes more precise when the contralateral inhibitory input propagates faster than the contralateral excitatory input. These results indicate that axon myelination and contralateral spike timing influence spatial hearing perception.