Codierung von Schallsignalen in Aktionspotenziale des auditorischen Nervs

Outer hair cells play a major role in the hearing process: they amplify the motion of the basilar membrane up to a 1000-fold and at the same time sharpen the excitation patterns. These patterns are converted by inner hair cells into action potentials of the auditory nerve. Outer hair cells are delicate structures and easily damaged, e. g., by overexposure to noise. Hearing aids can amplify the amplitude of the excitation patterns, but they cannot restore their degraded frequency selectivity. Noise overexposure also leads to delayed degeneration of auditory nerve fibers, particularly those with low a spontaneous rate, which are important for the coding of sound in noise. However, this loss cannot be diagnosed by pure-tone audiometry.     

Auditory Processing of Spectral Cues for Sound Localization in the Inferior Colliculus

The head-related transfer function (HRTF) of the cat adds directionally dependent energy minima to the amplitude spectrum of complex sounds. These spectral notches are a principal cue for the localization of sound source elevation. Physiological evidence suggests that the dorsal cochlear nucleus (DCN) plays a critical role in the brainstem processing of this directional feature. Type O units in the central nucleus of the inferior colliculus (ICC) are a primary target of ascending DCN projections and, therefore, may represent midbrain specializations for the auditory processing of spectral cues for sound localization. Behavioral studies confirm a loss of sound orientation accuracy when DCN projections to the inferior colliculus are surgically lesioned. This study used simple analogs of HRTF notches to characterize single-unit response patterns in the ICC of decerebrate cats that may contribute to the directional sensitivity of the brain's spectral processing pathways. Manipulations of notch frequency and bandwidth demonstrated frequency-specific excitatory responses that have the capacity to encode HRTF-based cues for sound source location. These response patterns were limited to type O units in the ICC and have not been observed for the projection neurons of the DCN. The unique spectral integration properties of type O units suggest that DCN influences are transformed into a more selective representation of sound source location by a local convergence of wideband excitatory and frequency-tuned inhibitory inputs.     

Subcortical neural coding mechanisms for auditory temporal processing

Biologically relevant sounds such as speech, animal vocalizations and music have distinguishing temporal features that are utilized for effective auditory perception. Common temporal features include sound envelope fluctuations, often modeled in the laboratory by amplitude modulation (AM), and starts and stops in ongoing sounds, which are frequently approximated by hearing researchers as gaps between two sounds or are investigated in forward masking experiments. The auditory system has evolved many neural processing mechanisms for encoding important temporal features of sound. Due to rapid progress made in the field of auditory neuroscience in the past three decades, it is not possible to review all progress in this field in a single article. The goal of the present report is to focus on single-unit mechanisms in the mammalian brainstem auditory system for encoding AM and gaps as illustrative examples of how the system encodes key temporal features of sound. This report, following a systems analysis approach, starts with findings in the auditory nerve and proceeds centrally through the cochlear nucleus, superior olivary complex and inferior colliculus. Some general principles can be seen when reviewing this entire field. For example, as one ascends the central auditory system, a neural encoding shift occurs. An emphasis on synchronous responses for temporal coding exists in the auditory periphery, and more reliance on rate coding occurs as one moves centrally. In addition, for AM, modulation transfer functions become more bandpass as the sound level of the signal is raised, but become more lowpass in shape as background noise is added. In many cases, AM coding can actually increase in the presence of background noise. For gap processing or forward masking, coding for gaps changes from a decrease in spike firing rate for neurons of the peripheral auditory system that have sustained response patterns, to an increase in firing rate for more central neurons with transient responses. Lastly, for gaps and forward masking, as one ascends the auditory system, some suppression effects become quite long (echo suppression), and in some stimulus configurations enhancement to a second sound can take place.     

Acoustic Basis of Directional Acuity in Laboratory Mice

The acoustic basis of auditory spatial acuity was investigated in CBA/129 mice by relating patterns of behavioral errors to directional features of the head-related transfer function (HRTF). Behavioral performance was assessed by training the mice to lick a water spout during sound presentations from a “safe” location and to suppress the response during presentations from “warning” locations. Minimum audible angles (MAAs) were determined by delivering the safe and warning sounds from different locations in the inter-aural horizontal and median vertical planes. HRTFs were measured at the same locations by implanting a miniature microphone and recording the gain of sound energy near the ear drum relative to free field. Mice produced an average MAA of 31° when sound sources were located in the horizontal plane. Acoustic measures indicated that binaural inter-aural level differences (ILDs) and monaural spectral features of the HRTF change systematically with horizontal location and therefore may have contributed to the accuracy of behavioral performance. Subsequent manipulations of the auditory stimuli and the directional properties of the ear produced errors that suggest the mice primarily relied on ILD cues when discriminating changes in azimuth. The MAA increased beyond 80° when the importance of ILD cues was minimized by testing in the median vertical plane. Although acoustic measures demonstrated a less robust effect of vertical location on spectral features of the HRTF, this poor performance provides further evidence for the insensitivity to spectral cues that was noted during behavioral testing in the horizontal plane.     

Frequency selectivity of phase-locking of complex sounds in the auditory nerve of the rat

Frequency selectivity of single auditory nerve fibers in the auditory nerve of the rat was studied using pseudorandom noise as the stimulus. The noise was lowpass filtered ternary m-sequences. Period histograms of the discharges of single auditory nerve fibers, locked to the periodicity of the noise, were cross-correlated with one period of the noise to obtain estimates of the impulse response. These cross-correlograms were subsequently Fourier transformed to obtain estimates of the frequency transfer functions. Earlier results obtained using noise that was based on binary sequences as the stimulus showed a systematic dependence on stimulus intensity of the bandwidth and center frequency of the computer transfer functions. The results of the present study confirmed this dependence and showed that a linear model based upon first-order cross-correlations fit the histograms of response. It is concluded that phase-locked activity of single auditory nerve fibers accurately reproduces the half-wave rectified motion of the basilar membrane over a large range of sound intensities.     

Intensity coding and the dynamic range problem

The psychophysical data on intensity discrimination indicate that certain schemes are unlikely as general intensity codes at the level of the auditory nerve and indirectly suggest that the most likely code is one based upon the firing rates of frequency-localized groups of fibers. A detection-theory analysis of a rate-based intensity code indicates that information from very few fibers can, if the information is appropriately combined, account for psychophysical discrimination even at high intensities. This suggests that fibers with similar CFs can code intensity over a wide range and that complex spectra can be represented at the level of the auditory nerve by a rate-CF code over the dynamic range of hearing. The analysis also indicates, however, a substantial discrepancy between the psychophysical data on the dependence of discrimination thresholds on level and the predicted discrimination behavior of a representative population of auditory nerve fibers. Thus, if intensity coding is based on localized firing rate, this fundamental psychophysical behavior does not result solely from peripheral processes.     

Quantifying Envelope and Fine-Structure Coding in Auditory Nerve Responses to Chimaeric Speech

Any sound can be separated mathematically into a slowly varying envelope and rapidly varying fine-structure component. This property has motivated numerous perceptual studies to understand the relative importance of each component for speech and music perception. Specialized acoustic stimuli, such as auditory chimaeras with the envelope of one sound and fine structure of another have been used to separate the perceptual roles for envelope and fine structure. Cochlear narrowband filtering limits the ability to isolate fine structure from envelope; however, envelope recovery from fine structure has been difficult to evaluate physiologically. To evaluate envelope recovery at the output of the cochlea, neural cross-correlation coefficients were developed that quantify the similarity between two sets of spike-train responses. Shuffled auto- and cross-correlogram analyses were used to compute separate correlations for responses to envelope and fine structure based on both model and recorded spike trains from auditory nerve fibers. Previous correlogram analyses were extended to isolate envelope coding more effectively in auditory nerve fibers with low center frequencies, which are particularly important for speech coding. Recovered speech envelopes were present in both model and recorded responses to one- and 16-band speech fine-structure chimaeras and were significantly greater for the one-band case, consistent with perceptual studies. Model predictions suggest that cochlear recovered envelopes are reduced following sensorineural hearing loss due to broadened tuning associated with outer-hair cell dysfunction. In addition to the within-fiber cross-stimulus cases considered here, these neural cross-correlation coefficients can also be used to evaluate spatiotemporal coding by applying them to cross-fiber within-stimulus conditions. Thus, these neural metrics can be used to quantitatively evaluate a wide range of perceptually significant temporal coding issues relevant to normal and impaired hearing.     

Representation of acoustic signals in the eighth nerve of the tokay gecko II. Masking of pure tones with noise

Acoustic signals are generally encoded in the peripheral auditory system of vertebrates by a duality scheme. For frequency components that fall within the excitatory tuning curve, individual eighth nerve fibers can encode the effective spectral energy by a spike-rate code, while simultaneously preserving the signal waveform periodicity of lower frequency components by phase-locked spike-train discharges. To explore how robust this duality of representation may be in the presence of noise, we recorded the responses of auditory fibers in the eighth nerve of the Tokay gecko to tonal stimuli when masking noise was added simultaneously. We found that their spike-rate functions reached plateau levels fairly rapidly in the presence of noise, so the ability to signal the presence of a tone by a concomitant change in firing rate was quickly lost. On the other hand, their synchronization functions maintained a high degree of phase-locked firings to the tone even in the presence of high-intensity masking noise, thus enabling a robust detection of the tonal signal. Critical ratios (CR) and critical bandwidths showed that in the frequency range where units are able to phaselock to the tonal periodicity, the CR bands were relatively narrow and the bandwidths were independent of noise level. However, to higher frequency tones where phaselocking fails and only spike-rate codes apply, the CR bands were much wider and depended upon noise level, so that their ability to filter tones out of a noisy background degraded with increasing noise levels. The greater robustness of phase-locked temporal encoding contrasted with spike-rate coding verifies a important advantage in using lower frequency signals for communication in noisy environments.     

Reversible long-term changes in auditory processing in mature auditory cortex in the absence of hearing loss induced by passive, moderate-level sound exposure

It has become increasingly clear that even occasional exposure to loud sounds in occupational or recreational settings can cause irreversible damage to the hair cells of the cochlea and the auditory nerve fibers, even if the resulting partial loss of hearing sensitivity, usually accompanied by tinnitus, disappears within hours or days of the exposure. Such exposure may explain at least some cases of poor speech intelligibility in noise in the face of a normal or near-normal audiogram. Recent findings from our laboratory suggest that long-term changes to auditory brain function-potentially leading to problems with speech intelligibility-can be effected by persistent, passive exposure to more moderate levels of noise (in the 70 dB SPL range) in the apparent absence of damage to the auditory periphery (as reflected in normal distortion product otoacoustic emissions and auditory brainstem responses). Specifically, passive exposure of adult cats to moderate levels of band-pass-filtered noise, or to band-limited ensembles of dense, random tone pips, can lead to a profound decrease of neural activity in the auditory cortex roughly in the exposure frequency range, and to an increase of activity outside that range. This can progress to an apparent reorganization of the cortical tonotopic map, which is reminiscent of the reorganization resulting from hearing loss restricted to a part of the hearing frequency range, although again, no hearing loss was apparent after our moderate-level sound exposure. Here, we review this work focusing specifically on the potential hearing problems that may arise despite a normally functioning auditory periphery.     

Advantages of directional hearing aid microphones related to room acoustics.

In this study, two types of hearing aids were used. Both aids had the same frequency characteristics for frontal sound, but one employed an omnidirectional microphone and the other a directional microphone. The frequency characteristics of both hearing aids were measured for five azimuths on KEMAR and in situ in 12 normal-hearing subjects. For these subjects we also determined the speech reception threshold (SRT) with background noise in two rooms with different reverberation times. The direction of the speech stimuli was always frontal; the direction of the noise was varied. Additionally, directional hearing was measured with short noise bursts from eight loudspeakers surrounding the subject. In the less reverberant room, sounds coming from behind were less amplified by the hearing aid with the directional microphone than by the one with the omnidirectional microphone. In this room the monaural SRT values were largely determined by the level of the background noise. For the directional hearing aids there was an extra binaural advantage which depended on the direction of the background noise. Only for low-frequency noise bursts was directional hearing better with directional hearing aids. In the more reverberant room, no distinct differences between the frequency characteristics of the two hearing aid types were measured. However, a systematic difference between monaural SRT values measured through the two hearing aids was found. This difference was independent of noise azimuth. In conclusion, hearing aid(s) with a directional microphone showed no disadvantages and clear advantages under specific conditions.     

Behavioral studies of the olivocochlear efferent system: learning to listen in noise

BACKGROUND: Olivocochlear (OC) neurons make up an efferent, descending auditory system that returns sound representations to the inner ear soon after they have entered the brain. Efferent inputs into the cochlea modulate outer hair cell activity to improve the neural encoding of auditory signals in background noise. Based on this physiological evidence, loss of efferent feedback is expected to degrade perception in noise. Attempts to confirm this prediction with long-term audiological assessments have met with mixed results. OBJECTIVE: To isolate procedural factors that may diminish the demonstration of long-term OC deficits in listening tasks. DESIGN: Operant conditioning procedures were used to train domestic cats to signal a change in the location of an auditory stimulus by responding on a lever. The smallest detectable change in location was measured by manipulating the distance between speakers under quiet conditions and in the presence of background noise. Functional consequences of efferent feedback were evaluated by comparing the sound localization thresholds of OC-lesioned cats with normal controls. RESULTS: As predicted by the hypothesized function of OC feedback systems, the lesioned cats exhibited significantly elevated thresholds only when tested in background noise. This initially poor performance returned to normal values after long-term exposure to the testing procedure. CONCLUSIONS: The results of our animal studies support the OC enhancement of sound localization behavior in background noise. Also, our behavioral observations suggest the acquisition of alternate listening strategies that allowed lesioned cats to minimize the functional consequences of their auditory deficits by attending more closely to remaining directional cues. These learned compensatory behaviors were encouraged by our present experimental design, which incorporated long-term training under consistent stimulus conditions. These findings point out the potential limitations of the highly routine audiological procedures that have been used to assess the impact of OC feedback on human hearing.     

Relationship between the auditory brainstem response and auditory nerve thresholds in cats with hearing loss

This study explored the relationship between the auditory brainstem response (ABR) and auditory nerve sensitivity in cats with normal hearing and with noise-induced permanent threshold shifts. A statistically significant linear correlation was found between each cat's ABR thresholds and the most sensitive single neuron thresholds at the same frequency. ABR thresholds were approximately 25 dB higher than the thresholds of the most sensitive neural responses in cats with normal hearing. The two measures produced equivalent thresholds at impaired frequencies in subjects with sensorineural hearing loss. Two factors may have contributed to this convergence of ABR and neural thresholds. First, our results suggest that the elevation of the most sensitive neural responses led to a compressed threshold distribution. Consequently, only a narrow range of sound levels separated stimulus conditions that activated relatively few fibers from those that were sufficient to evoke a robust population response. In addition, the threshold responses of impaired auditory nerve fibers may have been augmented by activity in the more sensitive 'off-frequency' regions that surrounded a discrete cochlear lesion. Across varying degrees of hearing loss, the ABR maintained a systematic relationship to auditory nerve fiber thresholds, and therefore has the potential to be used as a functional assay of cochlear pathology.     

Investigating the Effect of Cochlear Synaptopathy on Envelope Following Responses Using a Model of the Auditory Nerve

The healthy auditory system enables communication in challenging situations with high levels of background noise. Yet, despite normal sensitivity to pure tones, many listeners complain about having difficulties in such situations. Recent animal studies demonstrated that noise overexposure that produces temporary threshold shifts can cause the loss of auditory nerve (AN) fiber synapses (i.e., cochlear synaptopathy, CS), which appears to predominantly affect medium- and low-spontaneous rate (SR) fibers. In the present study, envelope following response (EFR) magnitude-level functions were recorded in normal hearing (NH) threshold and mildly hearing-impaired (HI) listeners with thresholds elevated above 2 kHz. EFRs were elicited by sinusoidally amplitude modulated (SAM) tones presented in quiet with a carrier frequency of 2 kHz, modulated at 93 Hz, and modulation depths of 0.85 (deep) and 0.25 (shallow). While EFR magnitude-level functions for deeply modulated tones were similar for all listeners, EFR magnitudes for shallowly modulated tones were reduced at medium stimulation levels in some NH threshold listeners and saturated in all HI listeners for the whole level range. A phenomenological model of the AN was used to investigate the extent to which hair-cell dysfunction and/or CS could explain the trends observed in the EFR data. Hair-cell dysfunction alone, including postulated elevated hearing thresholds at extended high frequencies (EHF) beyond 8 kHz, could not account for the recorded EFR data. Postulated CS led to simulations generally consistent with the recorded data, but a loss of all types of AN fibers was required within the model framework. The effects of off-frequency contributions (i.e., away from the characteristic place of the stimulus) and the differential loss of different AN fiber types on EFR magnitude-level functions were analyzed. When using SAM tones in quiet as the stimulus, model simulations suggested that (1) EFRs are dominated by the activity of high-SR fibers at all stimulus intensities, and (2) EFRs at medium-to-high stimulus levels are dominated by off-frequency contributions.

     

Hearing loss, disorders of sound localization with the preservation ofsmell and taste in diabetics

Diabetes is often complicated by serious medical conditions which could be related to the development of auditory system and cranial nerves lesions, disorder of sound localisation and decreased olfactory and taste ability. Cranial nerve palsies in diabetes are considered as an integral part of the main disease. Twenty nine children with diabetes and non-diabetics control group, without a history of exposure to noise, ototoxic drugs, or ear disease aged 4-19 years old were examined. Hearing impairment may be present in children with disease duration above 3-5 years and in children with other complications. Sound localisation tests were performed by the method of Zakrzewski from a free auditory field, measuring the angle of directional hearing acuity in 35 diabetics age from 16 to 78 years. Longer duration of diabetes was associated with higher directional hearing acuity. Higher hearing loss was observed with higher directional values in diabetics. The authors investigated smell and taste in 35 diabetics. All patients were treated with insulin. Impairment value of smell identification thresholds was much more frequent than impairment of smell perception. In no case raised threshold of taste perception.     

Midbrain Synchrony to Envelope Structure Supports Behavioral Sensitivity to Single-Formant Vowel-Like Sounds in Noise

Vowels make a strong contribution to speech perception under natural conditions. Vowels are encoded in the auditory nerve primarily through neural synchrony to temporal fine structure and to envelope fluctuations rather than through average discharge rate. Neural synchrony is thought to contribute less to vowel coding in central auditory nuclei, consistent with more limited synchronization to fine structure and the emergence of average-rate coding of envelope fluctuations. However, this hypothesis is largely unexplored, especially in background noise. The present study examined coding mechanisms at the level of the midbrain that support behavioral sensitivity to simple vowel-like sounds using neurophysiological recordings and matched behavioral experiments in the budgerigar. Stimuli were harmonic tone complexes with energy concentrated at one spectral peak, or formant frequency, presented in quiet and in noise. Behavioral thresholds for formant-frequency discrimination decreased with increasing amplitude of stimulus envelope fluctuations, increased in noise, and were similar between budgerigars and humans. Multiunit recordings in awake birds showed that the midbrain encodes vowel-like sounds both through response synchrony to envelope structure and through average rate. Whereas neural discrimination thresholds based on either coding scheme were sufficient to support behavioral thresholds in quiet, only synchrony-based neural thresholds could account for behavioral thresholds in background noise. These results reveal an incomplete transformation to average-rate coding of vowel-like sounds in the midbrain. Model simulations suggest that this transformation emerges due to modulation tuning, which is shared between birds and mammals. Furthermore, the results underscore the behavioral relevance of envelope synchrony in the midbrain for detection of small differences in vowel formant frequency under real-world listening conditions.     

Envelope Coding in Auditory Nerve Fibers Following Noise-Induced Hearing Loss

Recent perceptual studies suggest that listeners with sensorineural hearing loss (SNHL) have a reduced ability to use temporal fine-structure cues, whereas the effects of SNHL on temporal envelope cues are generally thought to be minimal. Several perceptual studies suggest that envelope coding may actually be enhanced following SNHL and that this effect may actually degrade listening in modulated maskers (e.g., competing talkers). The present study examined physiological effects of SNHL on envelope coding in auditory nerve (AN) fibers in relation to fine-structure coding. Responses were compared between anesthetized chinchillas with normal hearing and those with a mild–moderate noise-induced hearing loss. Temporal envelope coding of narrowband-modulated stimuli (sinusoidally amplitude-modulated tones and single-formant stimuli) was quantified with several neural metrics. The relative strength of envelope and fine-structure coding was compared using shuffled correlogram analyses. On average, the strength of envelope coding was enhanced in noise-exposed AN fibers. A high degree of enhanced envelope coding was observed in AN fibers with high thresholds and very steep rate-level functions, which were likely associated with severe outer and inner hair cell damage. Degradation in fine-structure coding was observed in that the transition between AN fibers coding primarily fine structure or envelope occurred at lower characteristic frequencies following SNHL. This relative fine-structure degradation occurred despite no degradation in the fundamental ability of AN fibers to encode fine structure and did not depend on reduced frequency selectivity. Overall, these data suggest the need to consider the relative effects of SNHL on envelope and fine-structure coding in evaluating perceptual deficits in temporal processing of complex stimuli.     

Modeling the Anti-masking Effects of the Olivocochlear Reflex in Auditory Nerve Responses to Tones in Sustained Noise

The medial olivocochlear reflex (MOCR) has been hypothesized to provide benefit for listening in noise. Strong physiological support for an anti-masking role for the MOCR has come from the observation that auditory nerve (AN) fibers exhibit reduced firing to sustained noise and increased sensitivity to tones when the MOCR is elicited. The present study extended a well-established computational model for normal-hearing and hearing-impaired AN responses to demonstrate that these anti-masking effects can be accounted for by reducing outer hair cell (OHC) gain, which is a primary effect of the MOCR. Tone responses in noise were examined systematically as a function of tone level, noise level, and OHC gain. Signal detection theory was used to predict detection and discrimination for different spontaneous rate fiber groups. Decreasing OHC gain decreased the sustained noise response and increased maximum discharge rate to the tone, thus modeling the ability of the MOCR to decompress AN fiber rate-level functions. Comparing the present modeling results with previous data from AN fibers in decerebrate cats suggests that the ipsilateral masking noise used in the physiological study may have elicited up to 20 dB of OHC gain reduction in addition to that inferred from the contralateral noise effects. Reducing OHC gain in the model also extended the dynamic range for discrimination over a wide range of background noise levels. For each masker level, an optimal OHC gain reduction was predicted (i.e., where maximum discrimination was achieved without increased detection threshold). These optimal gain reductions increased with masker level and were physiologically realistic. Thus, reducing OHC gain can improve tone-in-noise discrimination even though it may produce a “hearing loss” in quiet. Combining MOCR effects with the sensorineural hearing loss effects already captured by this computational AN model will be beneficial for exploring the implications of their interaction for the difficulties hearing-impaired listeners have in noisy situations.     

The representation of noise vocoded speech in the auditory nerve of the chinchilla: physiological correlates of the perception of spectrally reduced speech

This study investigated the neural representation of naturally produced and noise vocoded speech signals in the auditory nerve of the chinchilla. The syllables [see text] produced by male speakers were used to synthesize noise vocoded speech stimuli containing one, two, three and four bands of envelope modulated noise. The ensemble response of the auditory nerve, computed by pooling the PST histograms across many auditory nerve fibers, revealed temporal patterns in the responses to the natural tokens that uniquely identified the stop consonants. The responses to the 3- and 4-band noise vocoded tokens contained temporal patterns that were nearly identical to those observed for the natural tokens, while the responses to the 1- and 2-band tokens were significantly different (p<0.0001). The ALSR, ALIR and autocorrelation of the pooled PST histograms represented the detail of the frequency spectrum for a naturally produced vowel, while the driven rate was unreliable. Each of these spectral analyses failed to reveal significant information about the noise vocoded vowels. These results suggest that temporal patterns in the responses of the auditory nerve can provide the cues necessary for the recognition of noise vocoded stop consonants.     

Bilateral cochlear implants in long-term and short-term deafness

This case is of a 70-year-old man with single-sided deafness (SSD) in the right ear since childhood, who developed a sudden severe hearing loss in the left ear at the age of 63. Eventually, after he received cochlear implants in both ears, he started to present behavioural auditory processing skills associated with binaural hearing, such as improved ability to understand speech in the presence of background noise, and sound localization. Outcomes were measured using cortical auditory evoked potentials, speech perception in noise, sound localization tests, and a self-rating questionnaire. The results suggest that even after more than 50 years of unilateral deafness it was possible to develop binaural interaction and sound localization as a result of electric auditory stimulation.