Vera F. Prijs

Single-unit response at the round window of the guinea pig.

In guinea pigs the unit contribution (unit action potential, ap) to the response of the round window was computed following the method used by Kiang and co-workers (1976), i.e., fibre discharges registered by a microelectrode in the nerve were used as trigger pulses for the averaging process of the corresponding ap registered with a gross electrode at the round window. Normally the ap was independent of the fibre-CF, had a diphasic waveform, and its amplitude was about 0.1 microV. Small inter-animal differences were found in waveform and amplitude of the ap. In a pilot experiment exploring pathological influences on ap, the auditory nerve was stretched to mimic the effect of some acoustic nerve tumors. We found that the waveforms of both ap and compound action potential (CAP) changed. The results indicate that in normal guinea pig cochleas the existence of an elementary unit waveform can be used in the convolution of the CAP [Goldstein, M.H. Jr. and Kiang, N.Y.S. (1958) J. Acoust. Soc. Am. 30, 107-114]. In abnormal cochleas, however, deteriorated aps may disturb the simple convolution concept of the CAP.

Single-fibre and whole-nerve responses to clicks as a function of sound intensity in the guinea pig

This paper describes a study of the intensity dependence of click-evoked responses of auditory-nerve fibres in relation to the simultaneously recorded compound action potential (CAP). Condensation and rarefaction clicks were presented to normal hearing guinea pigs over an intensity range of 60 dB. The recorded poststimulus time histograms (PSTHs) were characterized by the latency (tp), amplitude (Ap) and synchronization (Sp) of their dominant peak, parameters that are particularly important for the understanding of the CAP. For all fibres tp decreased monotonically with increasing intensity, in a continuous way for fibres with high characteristic frequency (CF greater than 3 kHz), and in discrete steps of one CF-cycle for low-CF (CF less than or equal to 3 kHz) fibres. An additional analysis of PSTH envelopes revealed that average latency shifts with intensity are similar for all CFs above 2 kHz. For all fibres Ap increased monotonically with intensity; the increase was stronger and maximum values were larger for low-CF than for high-CF fibres. A schematic model PSTH was then formulated on the basis of the experimental data. A sum of these model PSTHs from a hypothesized fibre population was convolved with an elemental unit response (Versnel et al., 1992) in order to simulate the compound action potential. Synthesized CAPs agreed with experimental CAPs in their main aspects.

Narrow-band analysis of compound action potentials for several stimulus conditions in the guinea pig

Compound action potentials (AP) were recorded under various stimulus conditions in 31 guinea pigs. Stimulus attenuation, decrease of inter-stimulus interval, increase of the level of a continuous wide-band noise maker, and lowering the animals temperature all resulted in a drop of the AP amplitude and an increase in latency. A narrow-band analysis of the compound APs makes it possible to describe these AP changes in terms of the response behaviour of small cochlear regions according to their central frequencies. The results show that intensity-dependent changes in the AP parameters can be explained on the basis of the tuning properties of the auditory nerve fibres when the effect of the rise time of the tone-burst stimulus is taken into account. Shortening of the inter-stimulus interval produces a complex interaction in terms of tone-burst frequency and the region along the cochlear partition that contributes dominantly to the AP. It is concluded that response contributions from the narrow bands with a central frequency near the tone-burst frequency show the most adaptation. The change in amplitude for narrow-band responses under increased masking is similar to that for stimulus attenuation. It seems, however, that the underlying masking mechanism is more comparable to the adaptation mechanism. Cooling of the animal did not affect the sharpness of tuning. In all four recording situations there seems to be a decrease in the amount of synchronization of single-fibre responses as reflected in the width of narrow-band action potentials.

Group delays of distortion product otoacoustic emissions in the guinea pig.

This paper presents a comprehensive set of experimental data on group delays of distortion product otoacoustic emissions (DPOAEs) in the guinea pig. Group delays of the DPOAEs with frequencies 2f1-f2, 3f1-2f2, 4f1-3f2, and 2f2-f1 were measured with the phase gradient method. Both the f1- and the f2-sweep paradigm were used. Differences between the two sweep paradigms were investigated for the four DPOAEs, as well as the group delay differences between the DPOAEs. Analysis revealed larger group delays with the f2-sweep paradigm, but only for the lower sideband DPOAEs (with fdp < f1,f2). For the lower sideband cubic distortion product 2f1-f2, the f2-sweep delays were a factor of 1.17-1.54 larger than the f1-sweep delays, depending on frequency. The upper sideband DPOAE 2f2-f1 showed no significant difference between f1- and f2-sweep group delays, except for the highest and lowest f2 frequencies. Comparing the group delays of the DPOAEs for each sweep paradigm separately, equal group delays were found for all four DPOAEs measured with the f1-sweep. With the f2-sweep paradigm on the other hand, the group delays of the three lower sideband DPOAEs occurred to be larger than the group delays of the upper sideband DPOAE 2f2-f1. A tentative interpretation of the data in the context of proposed explanatory hypotheses on DPOAE group delays is given.

Lower boundaries of two-tone suppression regions in the guinea pig.

Lower boundaries of two-tone suppression regions were determined in single fibres of the guinea pig with a tracking algorithm as described by Schmiedt (1982). For a suppressee at CF having a level of 20 dB above the threshold of the tip, suppression at the high-frequency (hf) side of the FTC could almost always be found. With the method used, the percentage of fibres in which suppression could be found at the low-frequency (lf) side of the FTC decreased with decreasing CF. Moreover, the occurrence of lf-suppression decreased for lower suppressee levels for fibres with CF approximately 2-5 kHz. For each fibre the minimum level difference between lf-suppression boundary and tip threshold was larger than 20 dB, for the whole group of fibres the difference was 34 dB on average. The hf-suppression regions sometimes reached below the tip for fibres with CFs in the 4 kHz region. The frequency at the lowest level of the hf-suppression boundary, best suppression frequency or BSF, is related to the CF as: BSF = 0.55 + 1.13 CF. When the suppressee level increased, the lower boundary at the hf side shifted upwards with a rate greater than 1 dB/dB. On the whole the two-tone suppression data in the guinea pig agree with those found in other rodents.

Transmitter release in inner hair cell synapses: a model analysis of spontaneous and driven rate properties of cochlear nerve fibres

The inner hair cell (IHC) synapse is one of the stages of cochlear processing that determine the relation between sound pressure level and spike rate in auditory nerve fibres. Transmitter released in the non-stimulated condition is held responsible for the wide range of spontaneous spike rates (SR) observed in these fibres. Properties of stimulated spike activity in auditory nerve fibres, including rate threshold and operating range of a fibre, are known to systematically vary with SR. This paper presents a model analysis of the relation between IHC transmembrane potential and transmitter release rate as becoming manifest in these spontaneous and driven rate properties. A previously developed computational model is used to identify those transfer properties of its synapse section which lead to reproduction of the variation of rate thresholds, shapes of rate-intensity functions and maximal driven rate with SR known from the literature. First a simple additive release model, in which driven transmitter release depends linearly on IHC potential, is elaborated. Its results lead to the hypothesis that the true release function is non-linear and variable across synapses generating different SR. An exponential release function is then introduced, with parameters varying across SR in a physiologically dictated way. This approach leads to adequate reproduction of the variation in rate thresholds and rate-intensity functions with SR. Finally, the model is applied in an inverse way to directly estimate the release function from given rate-intensity functions. The conclusion of both forward and inverse model analyses is that transmitter release is a non-linear function of IHC potential which, by the systematic variation of its parameters across SR, effectively leads to the physiological variation in dynamic range across fibres of different SR. Possible relations of these results with ultrastructural morphology and basic physiology of IHC synapses are discussed.

Round-window recorded potential of single-fibre discharge (unit response) in normal and noise-damaged cochleas

Unit responses (URs) of eighth-nerve fibres have been determined at the round window by spike-triggered averaging in both normal and pathological guinea pig cochleas. The pathology was mainly noise-induced damage. The URs have been analysed with respect to their dependence on the fibres threshold, characteristic frequency (CF) and spontaneous rate (SR). The results from normal cochleas confirmed earlier data (Prijs, 1986): the UR has a diphasic waveform and the amplitude of its negative first peak is about 0.1 microV. From the six parameters (amplitude, latency, and width of the two peaks) by which the UR was described only the amplitude of the positive peak showed a significant variation with CF: a small decrease with increasing CF (CF-range 0.1 to 20 kHz). This finding may possibly be caused by oscillations in the spike-triggered average for low CFs. URs for most low- and medium-SR fibres were found to be large (greater than 0.3 microV). However, this result is interpreted as an artefact caused by synchrony of fibre spontaneous activity. In damaged cochleas only slight changes of the UR were found: the waveform duration became significantly shorter and on some occasions the positive peak increased in amplitude, but latency and amplitude of the negative component of the UR remained unchanged.

Recovery characteristics of auditory nerve fibres in the normal and noise-damaged guinea pig cochlea

Spontaneous activity was analysed in auditory-nerve fibres innervating normal and noise-damaged cochleas. Spike occurrences were conceived as point processes. Joint interval distributions and serial correlation coefficients reveal a weak history effect for succeeding intervals. The point process is regarded as a renewal and the recovery function, being proportional to the hazard function, is determined from the interval probability density function. In 29 out of 60 fibres the latter shows peculiarities which result in a deviation from a monotonically increasing recovery function. For three fibers of low characteristic frequency the interval probability function shows an oscillatory pattern and for 26 fibres this function exhibits an early, sharp peak around 1.1 ms irrespective of characteristic frequency, spontaneous rate, or cochlear damage. The recovery function is not different between fibres with normal and those with abnormally high thresholds and exhibits an exponential recovery with one time constant of average value 1.6 ms. Bursting activity is found in only one fibre from the abnormally high threshold group.

A dual filter model describing single‐fiber responses to clicks in the normal and noise‐damaged cochlea

This paper presents a composite model of the normal and noise-damaged guinea pig cochlea. The model incorporates a phenomenologically defined cochlear filter, and physiological descriptions of inner hair cell transduction, synaptic adaptation, and spike generation. The latter three model sections were taken from recent literature. The paper first deals with validation and evaluation of the model and adaptation of the relevant parameters to the guinea pig. Then the model is applied to explore to what extent changes in the cochlear filter can be held responsible for abnormal responses to clicks that were recorded in single auditory nerve fibers in noise-damaged animals. Focus is on those fibers in which the tip-to-tail sensitivity ratio of the frequency threshold curve (FTC) has decreased and/or in which the FTC tail has become hypersensitive. Inspired by this type of W-shaped FTC the mechanical response of the basilar membrane is phenomenologically modeled by two parallel filters, one responsible for the tip of the FTC, the other for its tail. Model simulations show that most abnormal temporal response properties can be explained by pathological alterations in the mechanical response. Residual discrepancies between model and experiment are identified which presumably point to pathological changes in other stages of cochlear processing.

Group delays of distortion product otoacoustic emissions: relating delays measured with f1- and f2-sweep paradigms.

A theoretical analysis is presented of group delays of distortion product otoacoustic emissions (DPOAEs) measured with the phase-gradient method. The aim of the analysis is to clarify the differences in group delays D1 and D2, obtained using the f1- and the f2-sweep paradigms, respectively, and the dependence of group delays on the order of the DPOAE. Two models are considered, the place-fixed and the wave-fixed models. While in the former model the generation place is assumed to be invariant with both f1- and f2-sweeps, in the latter model the shift of generation place is fully accounted for. By making a simple local approximation of the cochlear scale invariance, a mathematical conversion from phase-place to phase-frequency gradients is incorporated in the wave-fixed model. Under the assumption that the DPOAE (as recorded at the best f2/f1 ratio) is dominated by the contribution from the generation site and not by, e.g., reflection components, the analysis leads to simple expressions for the ratio and difference between D1 and D2. Validation of the models against experimental data indicates that lower sideband DPOAEs (2f1-f2, 3f1-2f2, 4f1-3f2) are most consistent with the wave-fixed model. Upper sideband components (2f2-f1), in contrast, are not properly described by either the place-fixed or the wave-fixed model, independent whether DPOAE generation is assumed to originate at the f2 or at the more basally located f(dp) characteristic place.