Longitudinal observations of the effects of ischemic stroke on binaural perception.

Acute ischemic stroke, characterized by a localized reduction in blood flow to specific areas of the brain, has been shown to affect binaural auditory perception. In a previous study conducted during the acute phase of ischemic stroke, two tasks of binaural hearing were performed: binaural tone-in-noise detection, and lateralization of stimuli with interaural time- or level differences. Various lesion-specific, as well as individual, differences in binaural performance between patients in the acute phase of stroke and a control group were demonstrated. For the current study, we re-invited the same group of patients, whereupon a subgroup repeated the experiments during the subacute and chronic phases of stroke. Similar to the initial study, this subgroup consisted of patients with lesions in different locations, including cortical and subcortical areas. At the group level, the results from the tone-in-noise detection experiment remained consistent across the three measurement phases, as did the number of deviations from normal performance in the lateralization task. However, the performance in the lateralization task exhibited variations over time among individual patients. Some patients demonstrated improvements in their lateralization abilities, indicating recovery, whereas others' lateralization performance deteriorated during the later stages of stroke. Notably, our analyses did not reveal consistent patterns for patients with similar lesion locations. These findings suggest that recovery processes are more individual than the acute effects of stroke on binaural perception. Individual impairments in binaural hearing abilities after the acute phase of ischemic stroke have been demonstrated and should therefore also be targeted in rehabilitation programs.

Fast binaural processing but sluggish masker representation reconfiguration.

Perceptual organization of complex acoustic scenes requires fast binaural processing for accurate localization or lateralization based on short single-source-dominated glimpses. This sensitivity also manifests in the ability to detect rapid oscillating interaural time and phase differences as well as interaural correlation. However, binaural processing has also been termed "sluggish" based on experiments that require binaural detection in a masker with an additional binaural cue change in temporal proximity. The present study shows that the temporal integration windows obtained from data on binaural sluggishness cannot account for the detection of rapid binaural oscillations. A model with fast IPD encoding but a slower process of updating the internal representation of the masker IPD statistics accounted for both experiments of the "fast" and the "sluggish" categories.

Characterization of the decline in auditory nerve phase locking at high frequencies.

The frequency dependence of phase locking in the auditory nerve influences various auditory coding mechanisms. The decline of phase locking with increasing frequency is commonly described by a low-pass filter. This study compares fitted low-pass filter parameters with the actual rate of phase locking decline. The decline is similar across studies and only 40 dB per decade, corresponding to the asymptotic decline of a second order filter.

Corrigendum: Effects of acute ischemic stroke on binaural perception.

[This corrects the article DOI: 10.3389/fnins.2022.1022354.].

Frequency dependence of sensitivity to interaural phase differences in pure tones.

It is well established that in normal-hearing humans, the threshold of interaural time differences for pure tones increases dramatically above about 1300 Hz, only to become unmeasurable above 1400 Hz. However, physiological data and auditory models suggest that the actual decline in sensitivity is more gradual and only appears to be abrupt because the maximum of the psychometric function dips below the threshold proportion correct, e.g., 0.794. Published data only report thresholds at certain proportions correct but not the decline of proportions correct or of the sensitivity index d' with increasing frequencies. Here, we present pure-tone behavioral data obtained with a constant stimulus procedure. Seven of nine subjects showed proportions correct above 0.9 at 1300 Hz and virtually no sensitivity at 1500 Hz (proportion correct within 0.07 of chance level). This corresponds to a sensitivity decline of 46-78 dB/oct, much steeper than predicted by existing models or by the decline of phase locking of the auditory nerve fibers in animal data.

Statistics of the instantaneous interaural parameters for dichotic tones in diotic noise (N0S ψ ).

Stimuli consisting of an interaurally phase-shifted tone in diotic noise-often referred to as N 0 S ψ -are commonly used to study binaural hearing. As a consequence of mixing diotic noise with a dichotic tone, this type of stimulus contains random fluctuations in both interaural phase- and level-difference. We report the joint probability density functions of the two interaural differences as a function of amplitude and interaural phase of the tone. Furthermore, a second joint probability density function for interaural phase differences and the instantaneous cross-power is derived. The closed-form expression can be used in future studies of binaural unmasking first to obtain the interaural statistics and then study more directly the relation between those statistics and binaural tone detection.

A hemispheric two-channel code accounts for binaural unmasking in humans.

Sound in noise is better detected or understood if target and masking sources originate from different locations. Mammalian physiology suggests that the neurocomputational process that underlies this binaural unmasking is based on two hemispheric channels that encode interaural differences in their relative neuronal activity. Here, we introduce a mathematical formulation of the two-channel model - the complex-valued correlation coefficient. We show that this formulation quantifies the amount of temporal fluctuations in interaural differences, which we suggest underlie binaural unmasking. We applied this model to an extensive library of psychoacoustic experiments, accounting for 98% of the variance across eight studies. Combining physiological plausibility with its success in explaining behavioral data, the proposed mechanism is a significant step towards a unified understanding of binaural unmasking and the encoding of interaural differences in general.

Lower interaural coherence in off-signal bands impairs binaural detection.

Differences in interaural phase configuration between a target and a masker can lead to substantial binaural unmasking. This effect is decreased for masking noises with an interaural time difference (ITD). Adding a second noise with an opposing ITD in most cases further reduces binaural unmasking. Thus far, modeling of these detection thresholds required both a mechanism for internal ITD compensation and an increased filter bandwidth. An alternative explanation for the reduction is that unmasking is impaired by the lower interaural coherence in off-frequency regions caused by the second masker [Marquardt and McAlpine (2009). J. Acoust. Soc. Am. 126(6), EL177-EL182]. Based on this hypothesis, the current work proposes a quantitative multi-channel model using monaurally derived peripheral filter bandwidths and an across-channel incoherence interference mechanism. This mechanism differs from wider filters since it has no effect when the masker coherence is constant across frequency bands. Combined with a monaural energy discrimination pathway, the model predicts the differences between a single delayed noise and two opposingly delayed noises as well as four other data sets. It helps resolve the inconsistency that simulating some data requires wide filters while others require narrow filters.

Considerations for Fitting Cochlear Implants Bimodally and to the Single-Sided Deaf.

When listening with a cochlear implant through one ear and acoustically through the other, binaural benefits and spatial hearing abilities are generally poorer than in other bilaterally stimulated configurations. With the working hypothesis that binaural neurons require interaurally matched inputs, we review causes for mismatch, their perceptual consequences, and experimental methods for mismatch measurements. The focus is on the three primary interaural dimensions of latency, frequency, and level. Often, the mismatch is not constant, but rather highly stimulus-dependent. We report on mismatch compensation strategies, taking into consideration the specific needs of the respective patient groups. Practical challenges typically faced by audiologists in the proposed fitting procedure are discussed. While improvement in certain areas (e.g., speaker localization) is definitely achievable, a more comprehensive mismatch compensation is a very ambitious endeavor. Even in the hypothetical ideal fitting case, performance is not expected to exceed that of a good bilateral cochlear implant user.

Simulation of ITD-Dependent Single-Neuron Responses Under Electrical Stimulation and with Amplitude-Modulated Acoustic Stimuli.

Interaural time difference (ITD) sensitivity with cochlear implant stimulation is remarkably similar to envelope ITD sensitivity using conventional acoustic stimulation. This holds true for human perception, as well as for neural response rates recorded in the inferior colliculus of several mammalian species. We hypothesize that robust excitatory-inhibitory (EI) interaction is the dominant mechanism. Therefore, we connected the same single EI-model neuron to either a model of the normal acoustic auditory periphery or to a model of the electrically stimulated auditory nerve. The model captured most features of the experimentally obtained response properties with electric stimulation, such as the shape of rate-ITD functions, the dependence on stimulation level, and the pulse rate or modulation-frequency dependence. Rate-ITD functions with high-rate, amplitude-modulated electric stimuli were very similar to their acoustic counterparts. Responses obtained with unmodulated electric pulse trains most resembled acoustic filtered clicks. The fairly rapid decline of ITD sensitivity at rates above 300 pulses or cycles per second is correctly simulated by the 3.1-ms time constant of the inhibitory post-synaptic conductance. As the model accounts for these basic properties, it is expected to help in understanding and quantifying the binaural hearing abilities with electric stimulation when integrated in bigger simulation frameworks.

Effects of acute ischemic stroke on binaural perception.

Stroke-induced lesions at different locations in the brain can affect various aspects of binaural hearing, including spatial perception. Previous studies found impairments in binaural hearing, especially in patients with temporal lobe tumors or lesions, but also resulting from lesions all along the auditory pathway from brainstem nuclei up to the auditory cortex. Currently, structural magnetic resonance imaging (MRI) is used in the clinical treatment routine of stroke patients. In combination with structural imaging, an analysis of binaural hearing enables a better understanding of hearing-related signaling pathways and of clinical disorders of binaural processing after a stroke. However, little data are currently available on binaural hearing in stroke patients, particularly for the acute phase of stroke. Here, we sought to address this gap in an exploratory study of patients in the acute phase of ischemic stroke. We conducted psychoacoustic measurements using two tasks of binaural hearing: binaural tone-in-noise detection, and lateralization of stimuli with interaural time- or level differences. The location of the stroke lesion was established by previously acquired MRI data. An additional general assessment included three-frequency audiometry, cognitive assessments, and depression screening. Fifty-five patients participated in the experiments, on average 5 days after their stroke onset. Patients whose lesions were in different locations were tested, including lesions in brainstem areas, basal ganglia, thalamus, temporal lobe, and other cortical and subcortical areas. Lateralization impairments were found in most patients with lesions within the auditory pathway. Lesioned areas at brainstem levels led to distortions of lateralization in both hemifields, thalamus lesions were correlated with a shift of the whole auditory space, whereas some cortical lesions predominantly affected the lateralization of stimuli contralateral to the lesion and resulted in more variable responses. Lateralization performance was also found to be affected by lesions of the right, but not the left, basal ganglia, as well as by lesions in non-auditory cortical areas. In general, altered lateralization was common in the stroke group. In contrast, deficits in tone-in-noise detection were relatively scarce in our sample of lesion patients, although a significant number of patients with multiple lesion sites were not able to complete the task.

Temporal hyper-precision of brainstem neurons alters spatial sensitivity of binaural auditory processing with cochlear implants.

The ability to localize a sound source in complex environments is essential for communication and navigation. Spatial hearing relies predominantly on the comparison of differences in the arrival time of sound between the two ears, the interaural time differences (ITDs). Hearing impairments are highly detrimental to sound localization. While cochlear implants (CIs) have been successful in restoring many crucial hearing capabilities, sound localization via ITD detection with bilateral CIs remains poor. The underlying reasons are not well understood. Neuronally, ITD sensitivity is generated by coincidence detection between excitatory and inhibitory inputs from the two ears performed by specialized brainstem neurons. Due to the lack of electrophysiological brainstem recordings during CI stimulation, it is unclear to what extent the apparent deficits are caused by the binaural comparator neurons or arise already on the input level. Here, we use a bottom-up approach to compare response features between electric and acoustic stimulation in an animal model of CI hearing. Conducting extracellular single neuron recordings in gerbils, we find severe hyper-precision and moderate hyper-entrainment of both the excitatory and inhibitory brainstem inputs to the binaural comparator neurons during electrical pulse-train stimulation. This finding establishes conclusively that the binaural processing stage must cope with highly altered input statistics during CI stimulation. To estimate the consequences of these effects on ITD sensitivity, we used a computational model of the auditory brainstem. After tuning the model parameters to match its response properties to our physiological data during either stimulation type, the model predicted that ITD sensitivity to electrical pulses is maintained even for the hyper-precise inputs. However, the model exhibits severely altered spatial sensitivity during electrical stimulation compared to acoustic: while resolution of ITDs near midline was increased, more lateralized adjacent source locations became inseparable. These results directly resemble recent findings in rodent and human CI listeners. Notably, decreasing the phase-locking precision of inputs during electrical stimulation recovered a wider range of separable ITDs. Together, our findings suggest that a central problem underlying the diminished ITD sensitivity in CI users might be the temporal hyper-precision of inputs to the binaural comparator stage induced by electrical stimulation.

Influence of envelope fluctuation on the lateralization of interaurally delayed low-frequency stimuli.

Disregarding onset and offset effects, interaurally delaying a 500 Hz tone by 1.5 ms is identical to advancing it by 0.5 ms. When presented over headphones, humans indeed perceive such a tone lateralized toward the side of the nominal lag. Any stimulus other than a tone has more than one frequency component and is thus unambiguous. It has been shown that phase ambiguity can be resolved when increasing the stimulus bandwidth. This has mostly been attributed to the integration of information across frequencies. Additionally, interaural timing information conveyed in the stimulus envelope within a single frequency channel is a second possible cue that could help to resolve phase ambiguity. This study employs stimuli designed to differ in the amount of envelope fluctuation while retaining the same power spectral density as well as interaural differences. Any difference in lateralization must thus be a result of the difference in envelope. The results show that stimuli with strong envelope fluctuation require significantly smaller bandwidths to resolve phase ambiguity when compared to stimuli with weak envelope fluctuation. This suggests that within-channel information is an important cue used to resolve phase ambiguity.

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.

Neural rate difference model can account for lateralization of high-frequency stimuli.

Lateralization of complex high-frequency sounds is conveyed by interaural level differences (ILDs) and interaural time differences (ITDs) in the envelope. In this work, the authors constructed an auditory model and simulate data from three previous behavioral studies obtained with, in total, over 1000 different amplitude-modulated stimuli. The authors combine a well-established auditory periphery model with a functional count-comparison model for binaural excitatory-inhibitory (EI) interaction. After parameter optimization of the EI-model stage, the hemispheric rate-difference between pairs of EI-model neurons relates linearly with the extent of laterality in human listeners. If a certain ILD and a certain envelope ITD each cause a similar extent of laterality, they also produce a similar rate difference in the same model neurons. After parameter optimization, the model accounts for 95.7% of the variance in the largest dataset, in which amplitude modulation depth, rate of modulation, modulation exponent, ILD, and envelope ITD were varied. The model also accounts for 83% of the variances in each of the other two datasets using the same EI model parameters.

Smallest perceivable interaural time differences.

It is well-established that the smallest discrimination thresholds for interaural time differences (ITDs) are near 10 µs for normal hearing listeners. However, little is known about the hearing and training status of the test subjects from past studies. Previous studies also did not explicitly focus on the identification of the optimal stimulus and measurement technique to obtain the smallest threshold ITDs. Therefore, the first goal of the current study was to identify the stimulus and experimental method that maximizes ITD sensitivity. The second goal was to provide a precise threshold ITD reference value for both well-trained and un-trained normal hearing listeners using the optimal stimulus and method. The stimulus that yielded the lowest threshold ITD was Gaussian noise, band-pass filtered from 20 to 1400 Hz, presented at 70 dB sound pressure level. The best method was a two-interval procedure with an interstimulus interval of 50 ms. The average threshold ITD for this condition at the 75% correct level was 6.9 µs for nine trained listeners and 18.1 µs for 52 un-trained listeners.

Coherent Coding of Enhanced Interaural Cues Improves Sound Localization in Noise With Bilateral Cochlear Implants.

Bilateral cochlear implant (BCI) users only have very limited spatial hearing abilities. Speech coding strategies transmit interaural level differences (ILDs) but in a distorted manner. Interaural time difference (ITD) information transmission is even more limited. With these cues, most BCI users can coarsely localize a single source in quiet, but performance quickly declines in the presence of other sound. This proof-of-concept study presents a novel signal processing algorithm specific for BCIs, with the aim to improve sound localization in noise. The core part of the BCI algorithm duplicates a monophonic electrode pulse pattern and applies quasistationary natural or artificial ITDs or ILDs based on the estimated direction of the dominant source. Three experiments were conducted to evaluate different algorithm variants: Experiment 1 tested if ITD transmission alone enables BCI subjects to lateralize speech. Results showed that six out of nine BCI subjects were able to lateralize intelligible speech in quiet solely based on ITDs. Experiments 2 and 3 assessed azimuthal angle discrimination in noise with natural or modified ILDs and ITDs. Angle discrimination for frontal locations was possible with all variants, including the pure ITD case, but for lateral reference angles, it was only possible with a linearized ILD mapping. Speech intelligibility in noise, limitations, and challenges of this interaural cue transmission approach are discussed alongside suggestions for modifying and further improving the BCI algorithm.

Better-ear glimpsing with symmetrically-placed interferers in bilateral cochlear implant users.

For a frontal target in spatially symmetrically placed interferers, normal hearing (NH) listeners can use "better-ear glimpsing" to select time-frequency segments with favorable signal-to-noise ratio in either ear. With an ideal monaural better-ear mask (IMBM) processing, some studies showed that NH listeners can reach similar performance as in the natural binaural listening condition, although interaural phase differences at low frequencies can further improve performance. In principle, bilateral cochlear implant (BiCI) listeners could use the same better-ear glimpsing, albeit without exploiting interaural phase differences. Speech reception thresholds of NH and BiCI listeners were measured in three interferers (speech-shaped stationary noise, nonsense speech, or single talker) either co-located with the target, symmetrically placed at ±60°, or independently presented to each ear, with and without IMBM processing. Furthermore, a bilateral noise vocoder based on the BiCI electrodogram was used in the same NH listeners. Headphone presentation and direct stimulation with head-related transfer functions for spatialization were used in NH and BiCI listeners, respectively. Compared to NH listeners, both NH listeners with vocoder and BiCI listeners showed strongly reduced binaural benefit from spatial separation. However, both groups greatly benefited from IMBM processing as part of the stimulation strategy.

Combination of binaural and harmonic masking release effects in the detection of a single component in complex tones.

Both harmonic and binaural signal properties are relevant for auditory processing. To investigate how these cues combine in the auditory system, detection thresholds for an 800-Hz tone masked by a diotic (i.e., identical between the ears) harmonic complex tone were measured in six normal-hearing subjects. The target tone was presented either diotically or with an interaural phase difference (IPD) of 180° and in either harmonic or "mistuned" relationship to the diotic masker. Three different maskers were used, a resolved and an unresolved complex tone (fundamental frequency: 160 and 40 Hz) with four components below and above the target frequency and a broadband unresolved complex tone with 12 additional components. The target IPD provided release from masking in most masker conditions, whereas mistuning led to a significant release from masking only in the diotic conditions with the resolved and the narrowband unresolved maskers. A significant effect of mistuning was neither found in the diotic condition with the wideband unresolved masker nor in any of the dichotic conditions. An auditory model with a single analysis frequency band and different binaural processing schemes was employed to predict the data of the unresolved masker conditions. Sensitivity to modulation cues was achieved by including an auditory-motivated modulation filter in the processing pathway. The predictions of the diotic data were in line with the experimental results and literature data in the narrowband condition, but not in the broadband condition, suggesting that across-frequency processing is involved in processing modulation information. The experimental and model results in the dichotic conditions show that the binaural processor cannot exploit modulation information in binaurally unmasked conditions.

A framework for testing and comparing binaural models.

Auditory research has a rich history of combining experimental evidence with computational simulations of auditory processing in order to deepen our theoretical understanding of how sound is processed in the ears and in the brain. Despite significant progress in the amount of detail and breadth covered by auditory models, for many components of the auditory pathway there are still different model approaches that are often not equivalent but rather in conflict with each other. Similarly, some experimental studies yield conflicting results which has led to controversies. This can be best resolved by a systematic comparison of multiple experimental data sets and model approaches. Binaural processing is a prominent example of how the development of quantitative theories can advance our understanding of the phenomena, but there remain several unresolved questions for which competing model approaches exist. This article discusses a number of current unresolved or disputed issues in binaural modelling, as well as some of the significant challenges in comparing binaural models with each other and with the experimental data. We introduce an auditory model framework, which we believe can become a useful infrastructure for resolving some of the current controversies. It operates models over the same paradigms that are used experimentally. The core of the proposed framework is an interface that connects three components irrespective of their underlying programming language: The experiment software, an auditory pathway model, and task-dependent decision stages called artificial observers that provide the same output format as the test subject.