Responses Of Auditory Nerve Fibers Research Articles

Biophysics-inspired spike rate adaptation for computationally efficient phenomenological nerve modeling

This study introduces and evaluates the PHAST+ model, part of a computational framework designed to simulate the behavior of auditory nerve fibers in response to the electrical stimulation from a cochlear implant. PHAST+ incorporates a highly efficient method for calculating accommodation and adaptation, making it particularly suited for simulations over extended stimulus durations. The proposed method uses a leaky integrator inspired by classic biophysical nerve models. Through evaluation against single-fiber animal data, our findings demonstrate the model’s effectiveness across various stimuli, including short pulse trains with variable amplitudes and rates. Notably, the PHAST+ model performs better than its predecessor, PHAST (a phenomenological model by van Gendt et al.), particularly in simulations of prolonged neural responses. While PHAST+ is optimized primarily on spike rate decay, it shows good behavior on several other neural measures, such as vector strength and degree of adaptation. The future implications of this research are promising. PHAST+ drastically reduces the computational burden to allow the real-time simulation of neural behavior over extended periods, opening the door to future simulations of psychophysical experiments and multi-electrode stimuli for evaluating novel speech-coding strategies for cochlear implants.

Simulating auditory nerve fiber response following micro-electrode stimulation

Abstract The cochlear implant was the first effective and is still the most common neuroprosthetic device which is employed for people with severe to profound hearing loss. To restore auditory perception, an array of micro-electrodes that deliver electrical pulses to the auditory nerve is surgically implanted into the lower cochlea duct, the scala tympani (ST). However, implantation into the upper cavity, the scala vestibuli (SV) has been tested due to severe anatomical obstruction or ossification of the ST. Clinical results revealed similar performance and thresholds for SV and ST cochlear implant users. We present a simulation study of auditory nerve fibre response to monophasic stimulation of both polarities. Excitation profiles are compared for microelectrodes placed in the SV and equivalent positions in the ST. In total, 7 different electrode positions for 4 different fibres have been investigated in a homogenous 2D model. Results for the intact fibres predict generally higher anodic thresholds in comparison to cathodic stimulation at the same electrode position and mostly lower thresholds for the SV electrodes in comparison to their ST counterparts. In contrast, anodic thresholds are mostly lower than cathodic thresholds for the degenerated fibres. Furthermore, due to the increased electrode-fibre distance for degenerate fibres which have completely lost the dendrite, SV stimulation is less beneficial. However, for basal fibres and the clinically relevant mid scala placement of the electrode the typically high thresholds remain similar for ST and SV positions.

Acoustically Evoked Compound Action Potentials Recorded From Cochlear Implant Users With Preserved Acoustic Hearing.

Less traumatic intracochlear electrode design and the introduction of the soft surgery technique allow for the preservation of low-frequency acoustic hearing in many cochlear implant (CI) users. Recently, new electrophysiologic methods have also been developed that allow acoustically evoked peripheral responses to be measured in vivo from an intracochlear electrode. These recordings provide clues to the status of peripheral auditory structures. Unfortunately, responses generated from the auditory nerve (auditory nerve neurophonic [ANN]) are somewhat difficult to record because they are smaller than the hair cell responses (cochlear microphonic). Additionally, it is difficult to completely segregate the ANN from the cochlear microphonic, complicating the interpretation and limiting clinical applications. The compound action potential (CAP) is a synchronous response of multiple auditory nerve fibers and may provide an alternative to ANN where the status of the auditory nerve is of primary interest. This study is a within-subject comparison of CAPs recorded using traditional stimuli (clicks and 500 Hz tone bursts) and a new stimulus (CAP chirp). We hypothesized that the chirp stimulus might result in a more robust CAP than that recorded using traditional stimuli, allowing for a more accurate assessment of the status of the auditory nerve. Nineteen adult Nucleus L24 Hybrid CI users with residual low-frequency hearing participated in this study. CAP responses were recorded from the most apical intracochlear electrode using a 100 μs click, 500 Hz tone bursts, and chirp stimuli presented via the insert phone to the implanted ear. The chirp stimulus used in this study was CAP chirp generated using parameters from human-derived band CAPs ( Chertoff et al. 2010 ). Additionally, nine custom chirps were created by systematically varying the frequency sweep rate of the power function used to construct the standard CAP chirp stimulus. CAPs were recorded using all acoustic stimuli, allowing for within-subject comparisons of the CAP amplitude, threshold, percentage of measurable CAP responses, and waveform morphology. Considerable variation in response morphology was apparent across stimuli and stimulation levels. Clicks and CAP chirp significantly evoked identifiable CAP response more compared to 500 Hz tone bursts. At relatively high stimulation levels, the chirp-evoked CAPs were significantly larger in amplitude and less ambiguous in morphology than the click-evoked CAPs. The status of residual acoustic hearing at high frequencies influenced the likelihood that a CAP could be reliably recorded. Subjects with better preserved hearing at high frequencies had significantly larger CAP amplitudes when CAP chirp was used. Customizing the chirp stimulus by varying the frequency sweep rates significantly affected the CAP amplitudes; however, pairwise comparisons did not show significant differences between chirps. CAPs can be measured more effectively using broadband acoustic stimuli than 500 Hz tone bursts in CI users with residual low-frequency acoustic hearing. The advantage of using CAP chirp stimulus relative to standard clicks is dependent on the extent of preserved acoustic hearing at high frequencies and the stimulus level. The chirp stimulus may present an attractive alternative to standard clicks or tone bursts for this CI population when the goal is to record robust CAP responses.

Human discrimination and modeling of high-frequency complex tones shed light on the neural codes for pitch.

Accurate pitch perception of harmonic complex tones is widely believed to rely on temporal fine structure information conveyed by the precise phase-locked responses of auditory-nerve fibers. However, accurate pitch perception remains possible even when spectrally resolved harmonics are presented at frequencies beyond the putative limits of neural phase locking, and it is unclear whether residual temporal information, or a coarser rate-place code, underlies this ability. We addressed this question by measuring human pitch discrimination at low and high frequencies for harmonic complex tones, presented either in isolation or in the presence of concurrent complex-tone maskers. We found that concurrent complex-tone maskers impaired performance at both low and high frequencies, although the impairment introduced by adding maskers at high frequencies relative to low frequencies differed between the tested masker types. We then combined simulated auditory-nerve responses to our stimuli with ideal-observer analysis to quantify the extent to which performance was limited by peripheral factors. We found that the worsening of both frequency discrimination and F0 discrimination at high frequencies could be well accounted for (in relative terms) by optimal decoding of all available information at the level of the auditory nerve. A Python package is provided to reproduce these results, and to simulate responses to acoustic stimuli from the three previously published models of the human auditory nerve used in our analyses.

Loss of central mineralocorticoid or glucocorticoid receptors impacts auditory nerve processing in the cochlea

SummaryThe key auditory signature that may associate peripheral hearing with central auditory cognitive defects remains elusive. Suggesting the involvement of stress receptors, we here deleted the mineralocorticoid and glucocorticoid receptors (MR and GR) using a CaMKIIα-based tamoxifen-inducible CreERT2/loxP approach to generate mice with single or double deletion of central but not cochlear MR and GR. Hearing thresholds of MRGRCaMKIIαCreERT2 conditional knockouts (cKO) were unchanged, whereas auditory nerve fiber (ANF) responses were larger and faster and auditory steady state responses were improved. Subsequent analysis of single MR or GR cKO revealed discrete roles for both, central MR and GR on cochlear functions. Limbic MR deletion reduced inner hair cell (IHC) ribbon numbers and ANF responses. In contrast, GR deletion shortened the latency and improved the synchronization to amplitude-modulated tones without affecting IHC ribbon numbers. These findings imply that stress hormone-dependent functions of central MR/GR contribute to “precognitive” sound processing in the cochlea.

Testing the plausibility of harmonic templates based on temporal fine-structure

The existence of harmonic templates in the central nervous system is a basic tenet of place models of pitch. Shamma & Klein (2000) propose that coincidences in temporal fine-structure across auditory nerve fibers in response to broadband stimuli suffice to set up harmonic templates for all fundamental frequencies in the phase-locking range. However, their computational model uses cochlear filters with unphysiologically steep slopes, calling into question the plausibility of the proposed scheme. We obtained responses from chinchilla auditory nerve fibers and “high-sync” neurons to pure tones, low-pass noise, and broadband noise, including “warped” noise (Heinz, 2006; Cedolin & Delgutte, 2010). Coincidences were computed using shuffled auto- or crosscorrelograms. We find that the maximum number of coincidences in response to tones of different frequencies, simulating the output of fibers with infinite sharp filters, indeed shows the template structure modeled by Shamma and Klein. This was not the case for responses to low-pass or broadband noise. However, if coincidences were only counted at zero delay, higher numbers of coincidences were indeed found for harmonically-related characteristic frequencies, even in response to noise. We conclude that steep-sloped cochlear filtering is not a necessary condition to obtain harmonic templates based on fine-structure.

Temporal Correlates to Monaural Edge Pitch in the Distribution of Interspike Interval Statistics in the Auditory Nerve.

Pitch is a perceptual attribute enabling perception of melody. There is no consensus regarding the fundamental nature of pitch and its underlying neural code. A stimulus which has received much interest in psychophysical and computational studies is noise with a sharp spectral edge. High-pass (HP) or low-pass (LP) noise gives rise to a pitch near the edge frequency (monaural edge pitch; MEP). The simplicity of this stimulus, combined with its spectral and autocorrelation properties, make it an interesting stimulus to examine spectral versus temporal cues that could underly its pitch. We recorded responses of single auditory nerve (AN) fibers in chinchilla to MEP-stimuli varying in edge frequency. Temporal cues were examined with shuffled autocorrelogram (SAC) analysis. Correspondence between the population’s dominant interspike interval and reported pitch estimates was poor. A fuller analysis of the population interspike interval distribution, which incorporates not only the dominant but all intervals, results in good matches with behavioral results, but not for the entire range of edge frequencies that generates pitch. Finally, we also examined temporal structure over a slower time scale, intermediate between average firing rate and interspike intervals, by studying the SAC envelope. We found that, in response to a given MEP stimulus, this feature also systematically varies with edge frequency, across fibers with different characteristic frequency (CF). Because neural mechanisms to extract envelope cues are well established, and because this cue is not limited by coding of stimulus fine-structure, this newly identified slower temporal cue is a more plausible basis for pitch than cues based on fine-structure.

Phenomenological model of auditory nerve population responses to cochlear implant stimulation

BackgroundModels of auditory nerve fiber (ANF) responses to electrical stimulation are helpful to develop advanced coding for cochlear implants (CIs). A phenomenological model of ANF population responses to CI electrical stimulation with a lower computational complexity compared to a biophysical model would be beneficial to evaluate new CI coding strategies. New methodThis study presents a phenomenological model which combines four temporal characteristics of ANFs (refractoriness, facilitation, accommodation and spike rate adaptation) in addition to a spatial spread of the electric field. ResultsThe model predicts the performances of CI subjects in the melodic contour identification (MCI) experiment. The simulations for the MCI experiment were consistent with CI recipients’ experimental outcomes that were not predictable from the electrical stimulation patterns themselves. Comparison with existing methodsPreviously, no phenomenological population model of ANFs has combined all four aforementioned temporal phenomena. ConclusionsThe proposed model would help the further investigations of ANFs responses to different electrical stimulation patterns and comparison of different sound coding strategies in CIs.

Cues to monaural edge pitch in single auditory nerve fibers

The spectral edge of a bandlimited noise can create a pitch sensation, called “edge pitch,” which is slightly mismatched to the edge frequency. This pitch is usually speculated to reflect neural lateral inhibition but could also reflect temporal cues. We looked for such cues recording responses of single auditory nerve fibers in the anesthetized chinchilla to edge pitch stimuli. For every stimulus, we compute shuffled autocorrelograms for all recorded fibers and pool them in a population interval distribution (PID) from which we obtain the most common interval. We also quantify the slower “envelope” fluctuations in firing resulting from cochlear bandpass filtering of the bandlimited noise stimulus. Single fibers shows systematic temporal changes related to the spectral edge of the stimulus when it does not cover the fiber's frequency filter. Across the population of fibers, the estimated pitch based on the dominant interspike interval shows trends that are similar, qualitatively and quantitatively, to those observed behaviorally. Slower envelope fluctuations also systematically vary with the relationship of edge frequency to fiber CF, and are another potential temporal cue to the presence of the edge frequency. We conclude that temporal cues to edge-pitch are present in the auditory nerve at multiple time scales.

Temporal Coding of Single Auditory Nerve Fibers Is Not Degraded in Aging Gerbils.

People suffering from age-related hearing loss typically present with deficits in temporal processing tasks. Temporal processing deficits have also been shown in single-unit studies at the level of the auditory brainstem, midbrain, and cortex of aged animals. In this study, we explored whether temporal coding is already affected at the level of the input to the central auditory system. Single-unit auditory nerve fiber recordings were obtained from 41 Mongolian gerbils of either sex, divided between young, middle-aged, and old gerbils. Temporal coding quality was evaluated as vector strength in response to tones at best frequency, and by constructing shuffled and cross-stimulus autocorrelograms, and reverse correlations, from responses to 1 s noise bursts at 10-30 dB sensation level (dB above threshold). At comparable sensation levels, all measures showed that temporal coding was not altered in auditory nerve fibers of aging gerbils. Furthermore, both temporal fine structure and envelope coding remained unaffected. However, spontaneous rates were decreased in aging gerbils. Importantly, despite elevated pure tone thresholds, the frequency tuning of auditory nerve fibers was not affected. These results suggest that age-related temporal coding deficits arise more centrally, possibly due to a loss of auditory nerve fibers (or their peripheral synapses) but not due to qualitative changes in the responses of remaining auditory nerve fibers. The reduced spontaneous rate and elevated thresholds, but normal frequency tuning, of aged auditory nerve fibers can be explained by the well known reduction of endocochlear potential due to strial dysfunction in aged gerbils.SIGNIFICANCE STATEMENT As our society ages, age-related hearing deficits become ever more prevalent. Apart from decreased hearing sensitivity, elderly people often suffer from a reduced ability to communicate in daily settings, which is thought to be caused by known age-related deficits in auditory temporal processing. The current study demonstrated, using several different stimuli and analysis techniques, that these putative temporal processing deficits are not apparent in responses of single-unit auditory nerve fibers of quiet-aged gerbils. This suggests that age-related temporal processing deficits may develop more central to the auditory nerve, possibly due to a reduced population of active auditory nerve fibers, which will be of importance for the development of treatments for age-related hearing disorders.

Comparison of Multi-Compartment Cable Models of Human Auditory Nerve Fibers.

Background: Multi-compartment cable models of auditory nerve fibers have been developed to assist in the improvement of cochlear implants. With the advancement of computational technology and the results obtained from in vivo and in vitro experiments, these models have evolved to incorporate a considerable degree of morphological and physiological details. They have also been combined with three-dimensional volume conduction models of the cochlea to simulate neural responses to electrical stimulation. However, no specific rules have been provided on choosing the appropriate cable model, and most models adopted in recent studies were chosen without a specific reason or by inheritance.Methods: Three of the most cited biophysical multi-compartment cable models of the human auditory nerve, i.e., Rattay et al. (2001b), Briaire and Frijns (2005), and Smit et al. (2010), were implemented in this study. Several properties of single fibers were compared among the three models, including threshold, conduction velocity, action potential shape, latency, refractory properties, as well as stochastic and temporal behaviors. Experimental results regarding these properties were also included as a reference for comparison.Results: For monophasic single-pulse stimulation, the ratio of anodic vs. cathodic thresholds in all models was within the experimental range despite a much larger ratio in the model by Briaire and Frijns. For biphasic pulse-train stimulation, thresholds as a function of both pulse rate and pulse duration differed between the models, but none matched the experimental observations even coarsely. Similarly, for all other properties including the conduction velocity, action potential shape, and latency, the models presented different outcomes and not all of them fell within the range observed in experiments.Conclusions: While all three models presented similar values in certain single fiber properties to those obtained in experiments, none matched all experimental observations satisfactorily. In particular, the adaptation and temporal integration behaviors were completely missing in all models. Further extensions and analyses are required to explain and simulate realistic auditory nerve fiber responses to electrical stimulation.

Divergent Auditory Nerve Encoding Deficits Between Two Common Etiologies of Sensorineural Hearing Loss.

Speech intelligibility can vary dramatically between individuals with similar clinically defined severity of hearing loss based on the audiogram. These perceptual differences, despite equal audiometric-threshold elevation, are often assumed to reflect central-processing variations. Here, we compared peripheral-processing in auditory nerve (AN) fibers of male chinchillas between two prevalent hearing loss etiologies: metabolic hearing loss (MHL) and noise-induced hearing loss (NIHL). MHL results from age-related reduction of the endocochlear potential due to atrophy of the stria vascularis. MHL in the present study was induced using furosemide, which provides a validated model of age-related MHL in young animals by reversibly inhibiting the endocochlear potential. Effects of MHL on peripheral processing were assessed using Wiener-kernel (system identification) analyses of single AN fiber responses to broadband noise, for direct comparison to previously published AN responses from animals with NIHL. Wiener-kernel analyses show that even mild NIHL causes grossly abnormal coding of low-frequency stimulus components. In contrast, for MHL the same abnormal coding was only observed with moderate to severe loss. For equal sensitivity loss, coding impairment was substantially less severe with MHL than with NIHL, probably due to greater preservation of the tip-to-tail ratio of cochlear frequency tuning with MHL compared with NIHL rather than different intrinsic AN properties. Differences in peripheral neural coding between these two pathologies-the more severe of which, NIHL, is preventable-likely contribute to individual speech perception differences. Our results underscore the need to minimize noise overexposure and for strategies to personalize diagnosis and treatment for individuals with sensorineural hearing loss.SIGNIFICANCE STATEMENT Differences in speech perception ability between individuals with similar clinically defined severity of hearing loss are often assumed to reflect central neural-processing differences. Here, we demonstrate for the first time that peripheral neural processing of complex sounds differs dramatically between the two most common etiologies of hearing loss. Greater processing impairment with noise-induced compared with an age-related (metabolic) hearing loss etiology may explain heightened speech perception difficulties in people overexposed to loud environments. These results highlight the need for public policies to prevent noise-induced hearing loss, an entirely avoidable hearing loss etiology, and for personalized strategies to diagnose and treat sensorineural hearing loss.

A Hybrid Learning Approach for Adaptive Classification of Acoustic Signals Using the Simulated Responses of Auditory Nerve Fibers

In this paper a framework of the heterogeneous system for an adaptive classification of multivariate numerical data is proposed. Considered data characterize the dynamics of the simulated responses of auditory nerve fibers as probability that represents the average of the ensemble for acoustic signals. The possibility to classify sound stimuli by means of analysis of the responses of the auditory nerve for the fibers with high and low spontaneous rates is examined. The aim of the study was to develop and implement a method for an adaptive pattern recognition in limited a priori information about their number and structure. The proposed model architecture consists of several units that generalize basic stages of a perceptual process that in turn corresponds neuro-symbolic information processing approach to the machine perception problem. Proposed method combines the advantages of the self-organizing maps and the radial basic function networks. This combination leads to a hybrid learning approach, which allow to provide the automatic classification of unlabeled data. According to the obtained results, the proposed approach showed better accuracy for several complex benchmark tests as well as for pure tones recognition by means of simulated auditory nerve fibers responses compared to k-means and single-linkage unsupervised classification strategies.

Modeling the effects of medial olivocochlear efferent stimulation at the level of the inferior colliculus.

Various studies on medial olivocochlear (MOC) efferents have implicated it in multiple roles in the auditory system (e.g., dynamic range adaptation, masking reduction, and selective attention). This study presents a systematic simulation of inferior colliculus (IC) responses with and without electrical stimulation of the MOC. Phenomenological models of the responses of auditory nerve (AN) fibers and IC neurons were used to this end. The simulated responses were highly consistent with physiological data (replicated 3 of the 4 known rate-level responses all MOC effects-shifts, high stimulus level reduction and enhancement). Complex MOC efferent effects which were previously thought to require integration from different characteristic frequency (CF) neurons were simulated using the same frequency inhibition excitation circuitry. MOC-induced enhancing effects were found only in neurons with a CF range from 750Hz to 2kHz. This limited effect is indicative of the role of MOC activation on the AN responses at the stimulus offset.

Phase Locking of Auditory-Nerve Fibers Reveals Stereotyped Distortions and an Exponential Transfer Function with a Level-Dependent Slope.

Phase locking of auditory-nerve-fiber (ANF) responses to the fine structure of acoustic stimuli is a hallmark of the auditory system's temporal precision and is important for many aspects of hearing. Period histograms from phase-locked ANF responses to low-frequency tones exhibit spike-rate and temporal asymmetries, but otherwise retain an approximately sinusoidal shape as stimulus level increases, even beyond the level at which the mean spike rate saturates. This is intriguing because apical cochlear mechanical vibrations show little compression, and mechanoelectrical transduction in the receptor cells is thought to obey a static sigmoidal nonlinearity, which might be expected to produce peak clipping at moderate and high stimulus levels. Here we analyze phase-locked responses of ANFs from cats of both sexes. We show that the lack of peak clipping is due neither to ANF refractoriness nor to spike-rate adaptation on time scales longer than the stimulus period. We demonstrate that the relationship between instantaneous pressure and instantaneous rate is well described by an exponential function whose slope decreases with increasing stimulus level. Relatively stereotyped harmonic distortions in the input to the exponential can account for the temporal asymmetry of the period histograms, including peak splitting. We show that the model accounts for published membrane-potential waveforms when assuming a power-of-three, but not a power-of-one, relationship to exocytosis. Finally, we demonstrate the relationship between the exponential transfer functions and the sigmoidal pseudotransducer functions obtained in the literature by plotting the maxima and minima of the voltage responses against the maxima and minima of the stimuli.SIGNIFICANCE STATEMENT Phase locking of auditory-nerve-fiber responses to the temporal fine structure of acoustic stimuli is important for many aspects of hearing, but the mechanisms underlying phase locking are not fully understood. Intriguingly, period histograms retain an approximately sinusoidal shape across sound levels, even when the mean rate has saturated. We find that neither refractoriness nor spike-rate adaptation is responsible for this behavior. Instead, the peripheral auditory system operates as though it contains an exponential transfer function whose slope changes with stimulus level. The underlying mechanism is distinct from the comparatively weak cochlear mechanical compression in the cochlear apex, and likely resides in the receptor cells.

Multidimensional stimulus encoding in the auditory nerve of the barn owl.

Auditory perception depends on multi-dimensional information in acoustic signals that must be encoded by auditory nerve fibers (ANF). These dimensions are represented by filters with different frequency selectivities. Multiple models have been suggested; however, the identification of relevant filters and type of interactions has been elusive, limiting progress in modeling the cochlear output. Spike-triggered covariance analysis of barn owl ANF responses was used to determine the number of relevant stimulus filters and estimate the nonlinearity that produces responses from filter outputs. This confirmed that ANF responses depend on multiple filters. The first, most dominant filter was the spike-triggered average, which was excitatory for all neurons. The second and third filters could be either suppressive or excitatory with center frequencies above or below that of the first filter. The nonlinear function mapping the first two filter outputs to the spiking probability ranged from restricted to nearly circular-symmetric, reflecting different modes of interaction between stimulus dimensions across the sample. This shows that stimulus encoding in ANFs of the barn owl is multidimensional and exhibits diversity over the population, suggesting that models must allow for variable numbers of filters and types of interactions between filters to describe how sound is encoded in ANFs.

Radon transform of auditory neurograms: a robust feature set for phoneme classification

Classification of speech phonemes is challenging, especially under noisy environments, and hence traditional speech recognition systems do not perform well in the presence of noise. Unlike traditional methods in which features are mostly extracted from the properties of the acoustic signal, this study proposes a new feature for phoneme classification using neural responses from a physiologically based computational model of the auditory periphery. The two-dimensional neurogram was constructed from the simulated responses of auditory-nerve fibres to speech phonemes. Features of neurogram images were extracted using the Discrete Radon Transform, and the dimensionality of features was reduced using an efficient feature selection technique. A standard classifier, Support Vector Machine, was employed to model and test the phoneme classes. Classification performance was evaluated in quiet and under noisy conditions in which test data were corrupted with various environmental distortions such as additive noise, room reverberation, and telephone-channel noise. Performances were also compared with the results from existing methods such as the Mel-frequency cepstral coefficient, Gammatone frequency cepstral coefficient, and frequency-domain linear prediction-based phoneme classification methods. In general, the proposed neural feature exhibited a better classification accuracy in quiet and under noisy conditions compared with the performance of most existing acoustic-signal-based methods.

Midbrain sensitivity to frequency “chirps:” Implications for coding voiced speech sounds

Many neurons in the mammalian midbrain (inferior colliculus, IC) have strong selectivity for direction of frequency transitions (“chirps”), which often co-exist with the better-known sensitivity for amplitude-modulation (AM) frequency. Chirps occur when harmonics with a phase gradient are summed, as in Schroeder-phase harmonic stimuli. Chirps also occur in voiced speech due to the phase properties of vocal tract resonances. IC neurons are direction-selective for chirps in stimuli with fundamental frequencies in the voice-pitch range, and chirp selectivity can be achieved in an AM-tuned IC model by adding off-CF inhibition. Here, we will show physiological and model responses illustrating how chirps influence IC responses to speech in awake rabbit. In healthy auditory-nerve (AN) responses, the frequency extent, or size, of chirps vary with proximity of AN tuning to formant peaks due to inner ear nonlinearities. For example, synchrony capture causes one harmonic to dominate responses of AN fibers near spectral peaks, and responses that are dominated by one harmonic do not exhibit chirps. The systematic variation in chirp size across frequency channels and strong selectivity of IC neurons for chirps suggest a role for this feature in coding voiced speech. Importantly, this coding mechanism is vulnerable to sensorineural hearing loss.

Measurements and models of auditory nerve spike rate and timing to electrical stimulation

Physiological measurements of the response of auditory nerve (AN) fibers to electrical stimulation from a cochlear implant (CI) have in general exhibited much higher maximal entrainment rates and higher temporal precision than is observed for acoustical stimulation. However, it appears that this superior rate coding in the AN for electrical stimulation does not translate to better coding of rate in the midbrain or for the percept of temporal pitch. Similarly, the enhanced temporal precision for electrical stimulation does not lead to improved coding of interaural timing differences. In this talk, I will review physiological data and computational model simulations that indicate that the superior spike rate and timing representation for CI stimulation may only hold when considering the response of some single AN fibers to repeated identical stimuli. In fact, there is a large heterogeneity in spiking responses that can change dramatically as a function of stimulus current level. A multi-compartmental computational model will be used to demonstrate how the electrode- fiber geometry and the membrane biophysics could be contributing to these phenomena. I will also discuss the implications of the likely population response of the AN to CI stimulation for brainstem processing. [Work supported by NSERC Discovery Grant 261736.]

Modeling the level-dependent changes of concurrent vowel scores.

The difference in fundamental frequency (F0) between talkers is an important cue for speaker segregation. To understand how this cue varies across sound level, Chintanpalli, Ahlstrom, and Dubno [(2014). J. Assoc. Res. Otolaryngol. 15, 823-837] collected level-dependent changes in concurrent-vowel identification scores for same- and different-F0 conditions in younger adults with normal hearing. Modeling suggested that level-dependent changes in phase locking of auditory-nerve (AN) fibers to formants and F0s may contribute to concurrent-vowel identification scores; however, identification scores were not predicted to test this suggestion directly. The current study predicts these identification scores using the temporal responses of a computational AN model and a modified version of Meddis and Hewitt's [(1992). J. Acoust. Soc. Am. 91, 233-245] F0-based segregation algorithm. The model successfully captured the level-dependent changes in identification scores of both vowels with and without F0 difference, as well as identification scores for one vowel correct. The model's F0-based vowel segregation was controlled using the actual F0-benefit across levels such that the predicted F0-benefit matched qualitatively with the actual F0-benefit as a function of level. The quantitative predictions from this F0-based segregation algorithm demonstrate that temporal responses of AN fibers to vowel formants and F0s can account for variations in identification scores across sound level and F0-difference conditions in a concurrent-vowel task.