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C D Geisler

Publications and source records attributed to C D Geisler.

At least 19 recordsLinked to original sources

Two-tone suppression by a saturating feedback model of the cochlear partition.

A model of a small strip of cochlear partition was computer simulated. The model is composed of two elements, approximations to the transfer functions of an inner hair cell (IHC) and an outer hair cell (OHC), respectively. The IHC element was insensitive to DC stimulation. Input was one or two sinusoids. One sinusoid, at the characteristic frequency (CF), was multiplied by the gain of the 'cochlear amplifier'. A second sinusoid, representing a tone with much lower frequency, was not affected by the amplifier gain. This gain was determined by the OHC transfer function. In one form of the model ('fixed-gain'), this gain was set at a fixed number determined from the furthest point reached on the OHC transfer function. This form of the model produced very realistic single-tone responses as well as showing 'two-tone suppression': that is, the IHC DC response produced by CF stimulation was reduced when the lower-frequency sinusoid, at suitable intensities, was added to the stimulus. When a DC component was added to the two-tone stimulus, the magnitude of this two-tone suppression was enhanced. In the second form of the model ('variable-gain'), the cochlear-amplifier gain varied throughout the stimulus cycle. Its value was re-calculated at each instant, determined by the point on the OHC transfer function current at that particular instant. This form of the model showed two-tone suppression only when a DC component was added to the two-tone stimulus.

Acoustic Stimulation

Two-tone suppression, excitation and the after effect in rate responses in auditory nerve fibres in the cat.

Responses were recorded from single, auditory nerve fibres in the anaesthetized cat. Acoustic stimuli consisted of two tones, one of which was at characteristic frequency (CF), the other (the suppressor) was at considerably lower frequency. Tones were presented in simultaneous and sequential configurations. For simultaneous presentations, well-known response properties were observed. The rising limb of the two-tone rate-intensity function closely matched that of the appropriately adapted response to the suppressor tone presented alone. Also, whether strongly suppressed relative to CF-driven rate, or equal to CF-driven rate, rate responses to the two-tone stimuli persisted unchanged when the CF tone was terminated and the suppressor tone continued alone. These results support the hypothesis that the suppressor tone has dual influences, suppressive and excitatory, that are distinct and additive. Peristimulus response histograms confirm in the cat that depression and slow recovery of sensitivity to CF may follow termination of the suppressor tone, as reported for the guinea pig [Hill, K.G. and Palmer, A.R. (1991) Hear. Res. 55, 167-176]. This delay in recovery of normal sensitivity to CF appeared to be directly related to the amount of excitation of the fibre that is attributable to the suppressor tone. A similar, delayed re-establishment of sensitivity also occurred in the response to a tone at CF, presented immediately following excitation by a suppressor tone. However, no delay occurred in the onset of response to the suppressor when preceded by the CF tone.(ABSTRACT TRUNCATED AT 250 WORDS)

Acoustic Stimulation

A cochlear model using feedback from motile outer hair cells.

A model of cochlear vibrations based upon motile outer hair cells (OHCs) has been developed using physiologically demonstrated phenomena. Rapid longitudinally directed OHC forces are connected in such a way as to form a negative-feedback system. The responses at the higher frequencies (greater than 1 kHZ) are quite realistic: they have properly shaped amplitude curves with large tip-to-tail ratios (30-50 dB), Q10's of 2-6, and 'shoulders' at frequencies an octave below the resonant frequency. The phases are also quite realistic, though asymptoting at somewhat lower values (about -6 pi radians) than observed physiologically. The responses in the apical section are not so realistic. The form of the OHC force is physically unrealizable, but realizable forms are discussed.

Acoustic Stimulation

Responses of "lower-spontaneous-rate" auditory-nerve fibers to speech syllables presented in noise. I: General characteristics.

Responses of auditory-nerve fibers in anesthetized cats to nine different spoken stop- and nasal-consonant/vowel syllables presented at 70 dB SPL in various levels of speech-shaped noise [signal-to-noise (S/N) ratios of 30, 20, 10, and 0 dB] are reported. The temporal aspects of speech encoding were analyzed using spectrograms. The responses of the "lower-spontaneous-rate" fibers (less than 20/s) were found to be more limited than those of the high-spontaneous-rate fibers. The lower-spontaneous-rate fibers did not encode noise-only portions of the stimulus at the lowest noise level (S/N = 30 dB) and only responded to the consonant if there was a formant or major spectral peak near its characteristic frequency. The fibers' responses at the higher noise levels were compared to those obtained at the lowest noise level using the covariance as a quantitative measure of signal degradation. The lower-spontaneous-rate fibers were found to preserve more of their initial temporal encoding than high-spontaneous-rate fibers of the same characteristic frequency. The auditory-nerve fibers' responses were also analyzed for rate-place encoding of the stimuli. The results are similar to those found for temporal encoding.

Animals

Responses of "lower-spontaneous-rate" auditory-nerve fibers to speech syllables presented in noise. II: Glottal-pulse periodicities.

The responses of single cat auditory-nerve fibers to naturally spoken voiced sounds (the vowels [a, i, u] and the murmur [m]) presented at normal intensity (70 dB SPL) in various levels of speech-shaped noise were analyzed for the encoding of the glottal-pulse (fundamental) period. To quantify the strength of this fundamental-period encoding, selected segments of the response histograms were autocorrelated, rectified, and fitted with the best-fitting sinusoid of the fundamental frequency. The magnitude of this best-fitting sinusoid was taken as the magnitude of synchronization. In most cases, it was found that the "lower-SR" fibers (those with spontaneous discharge rates less than 20/s) encoded the fundamental periodicity more strongly and more robustly than did the "high-SR" fibers (those with spontaneous discharge rates greater than 20/s). When either a single strong spectral peak or a relatively "flat" spectrum excited a fiber, it showed poor synchronization to the fundamental period, regardless of its spontaneous-rate class. Judging from a few examples, the glottal-pulse synchronization appears to be intensity dependent, with the relative performance of the high-SR fibers improving at lower intensities. A conceptual model is given which accounts for the general characteristics of the data.

Animals

Evidence for expansive power functions in the generation of the discharges of 'low- and medium-spontaneous' auditory-nerve fibers.

Re-analysis of data from Geisler et al. [J. Acoust. Soc. Am. 77, 1102-1109, 1985] indicates that the slopes of the intensity versus discharge-rate curves of auditory nerve (AN) fibers decrease systematically with increasing spontaneous discharge rate. For 'high-spontaneous' fibers, the slope is usually less than 0.5 dB/dB, while for 'low-spontaneous' fibers the slopes reach values greater than 4.0 dB/dB. A two-stage model accounts for this behavior. The first stage is a static non-linearity based on the measured intensity-voltage characteristic of inner hair cells. The second stage, representing action-potential generation, is linear for high-spontaneous fibers, but a squaring function for low- and medium-spontaneous fibers. The output of the model displays realistic slopes for its various intensity-rate curves. There are suggestions that a nonlinearity of still higher power is needed to simulate accurately the behavior of AN fibers having the lowest spontaneous rates (less than 0.1/s). The model also accounts for other observed differences between the discharge patterns of the different fiber classes.

Action Potentials

Saturation of outer hair cell receptor currents causes two-tone suppression.

Zwicker [Biol. Cybern. 35, 243-250, (1979); J. Acoust. Soc. Am. 80, 163-176 (1986)] has previously proposed that many nonlinear phenomena in the mammalian cochlea can be explained by saturation of a positive feedback process which enhances mechanical sensitivity, although the site of the nonlinearity producing this saturation has so far remained obscure. In this paper we present evidence suggesting that the nonlinearity of mechano-electrical transduction in the outer hair cells is the dominant nonlinearity producing two-tone suppression in the mammalian cochlea. In particular, we show that: (i) suppression of the extracellular summating potential (SP), recorded from a particular place within the organ of Corti, has characteristics similar to the suppression of activity in the auditory-nerve; (ii) that SP suppression occurs at approximately constant basilar membrane displacement, inferred from the SP iso-response contours; and that (iii) the onset of SP suppression with suppressor tones on the tail of the frequency tuning curve closely parallels the onset of nonlinearity in the local cochlear microphonic. Since previous studies (Patuzzi et al., 1989) have demonstrated that the vibration of the basilar membrane at its characteristic frequency is very sensitive to changes in outer hair cell receptor current, we consider that interference in outer hair cell currents caused by nonlinearity in mechano-electrical transduction is an adequate explanation of two-tone suppression. This requires that outer hair cell receptor currents deviate from linearity at a suppressor tone level below that required to produce a significant DC receptor potential within the inner hair cells, and that the active process within the cochlea is distributed along a local region of the cochlea, basal of the vibration peak.

Acoustic Stimulation

Estimation of eardrum acoustic pressure and of ear canal length from remote points in the canal.

Sound pressure distributions in the human ear canal, whether unoccluded or occluded with ear molds, were studied using a probe tube technique. On average, for frequencies below 6 kHz, the measuring probe tube had to be placed within 8 mm of the vertical plane containing the top of the eardrum (TOD), determined optically, in order to obtain sound pressure magnitudes within 6 dB of "eardrum pressure." To obtain that accuracy in all of the eight subjects studied, the probe had to be within 6 mm of the TOD. Since probe location relative to the drum has to be known, a purely acoustic method was developed which can be conveniently used to localize the probe-tip position, utilizing the standing wave property of the sound pressure in the ear canal. The acoustically estimated "drum location" generally lay between the optically determined vertical planes containing the TOD and the umbo. On average, the "drum location" fell 1 mm medial to the TOD. Of the 32 estimates made acoustically in various occluded and unoccluded conditions in 14 subjects, 30 estimates lay within a +/- 2-mm range of this average.

Acoustic Stimulation

Responses of "high-spontaneous" auditory-nerve fibers to consonant-vowel syllables in noise.

Responses of "high-spontaneous" single auditory-nerve fibers in anesthetized cat to nine different spoken stop and nasal consonant-vowel syllables presented in four different levels of speech-shaped noise are reported. The temporal information contained in the responses was analyzed using "composite" spectrograms and pseudo-3D spatial-frequency plots. Spectral characteristics of both consonant and vowel segments of the CV syllables were strongly encoded at S/N ratios of 30 and 20 dB. At S/N = 10 dB, formant information during the vowel segments was all that was reliably detectable in most cases. Even at S/N = 0 dB, most vowel formants were detectable, but only with relatively long analysis windows (40 ms). The increases (and decreases) in discharge rate during various phases of the responses were also determined. The rate responses to the "release" and to the voicing of the stop-consonant syllables were quite robust, being detectable at least half of the time, even at the highest noise level. Comparisons with psychoacoustic studies using similar stimuli are made.

Animals

The responses of models of "high-spontaneous" auditory-nerve fibers in a damaged cochlea to speech syllables in noise.

The responses of four high-spontaneous fibers from a damaged cat cochlea responding to naturally uttered consonant-vowel (CV) syllables [m], [p], and [t], each with [a], [i], and [u] in four different levels of noise were simulated using a two-stage computer model. At the lowest noise level [+30 dB signal-to-noise (S/N) ratio], the responses of the models of the three fibers from a heavily damaged portion of the cochlea [characteristic frequencies (CFs) from 1.6 to 2.14 kHz] showed quite different response patterns from those of fibers in normal cochleas: There was little response to the noise alone, the consonant portions of the syllables evoked small-amplitude wide-bandwidth complexes, and the vowel-segment response synchrony was often masked by low-frequency components, especially the first formant. At the next level of noise (S/N = 20 dB), spectral information regarding the murmur segments of the [m] syllables was essentially lost. At the highest noise levels used (S/N = +10 and 0 dB), the noise was almost totally disruptive of coding of the spectral peaks of the consonant portions of the stop CVs. Possible implications of the results with regard to the understanding of speech by hearing-impaired listeners are discussed.

Acoustic Stimulation

Responses of auditory-nerve fibers to nasal consonant-vowel syllables.

Responses of single auditory-nerve fibers in anesthetized cat to spoken nasal consonant-vowel syllables were recorded. Analyses in the form of spectrograms and of three-dimensional spatial-time and spatial-frequency plots were made. Among other features, formant transitions are clearly represented in the fibers' response synchronization properties. During vocalic segments, especially those in /mu/and/ma/, at a stimulus level near 75 dB SPL, a strong dominance in the responses by frequencies near the second formant (F2) is found for most fibers whose characteristic frequencies (CFs) are at or above F2. In contrast, at more moderate levels, the same fibers may show response synchrony to frequencies closer to their own CFs. There are significant differences in the response properties of high and low/medium-spontaneous-rate fibers.

Animals

Responses of auditory-nerve fibers to multiple-tone complexes.

To relate level-dependent properties of auditory-nerve-fiber responses to nasal consonant-vowels to the basic frequency selective and suppressive properties of the fibers, multitone complexes, with the amplitude of a single (probe) component incremented, were used as stimuli. Quantitative relations were obtained between the systematic increase of fiber synchrony to the probe tone and the decrease of synchrony to CF, as the amplitude of the probe tone was increased. When such relations are interpreted as a measure of fiber frequency selectivity based on a relative synchrony criterion, a breadth of frequency tuning is obtained, at a 70-dB SPL multitone sound-pressure level, which is generally broader than that of the fiber's threshold tuning curve. Quantitative comparisons with the same fiber's responses to the nasal speech sounds indicate that the fiber's speech responses share some common features with its probe-tone responses.

Animals

A composite auditory model for processing speech sounds.

A composite inner-ear model, containing the middle ear, basilar membrane (BM), hair cells, and hair-cell/nerve-fiber synapses, is presented. The model incorporates either a linear-BM stage or a nonlinear one. The model with the nonlinear BM generally shows a high degree of success in reproducing the qualitative aspects of experimentally recorded cat auditory-nerve-fiber responses to speech. In modeling fiber population responses to speech and speech in noise, it was found that the BM nonlinearity allows bands of fibers in the model to synchronize strongly to a common spectral peak in the stimulus. A cross-channel correlation algorithm has been devised to further process the model's population outputs. With output from the nonlinear-BM model, the cross-channel correlation values are appreciably reduced only at those channels whose CFs coincide with the formant frequencies. This observation also holds, to a large extent, for noisy speech.

Basilar Membrane

A model of the effect of outer hair cell motility on cochlear vibrations.

A model of cochlear function is presented in which deformation forces within outer hair cells are assumed to occur in synchronized response to generator potentials. Assuming a 90 degree phase lag between the generator potentials and the deformation forces, it is shown that the forces act to reduce cochlear-partition damping and thus increase frequency selectivity. A number of other experimentally observed phenomena, such as the effects of efferent-fiber stimulation and electrical polarization, can also be accounted for with this model.

Basilar Membrane

A temporal analysis of auditory-nerve fiber responses to spoken stop consonant-vowel syllables.

Auditory-nerve fiber spike trains were recorded in response to spoken English stop consonant-vowel syllables, both voiced (/b,d,g/) and unvoiced (/p,t,k/), in the initial position of syllables with the vowels /i,a,u/. Temporal properties of the neural responses and stimulus spectra are displayed in a spectrographic format. The responses were categorized in terms of the fibers' characteristic frequencies (CF) and spontaneous rates (SR). High-CF, high-SR fibers generally synchronize to formants throughout the syllables. High-CF, low/medium-SR fibers may also synchronize to formants; however, during the voicing, there may be sufficient low-frequency energy present to suppress a fiber's synchronized response to a formant near its CF. Low-CF fibers, from both SR groups, synchronize to energy associated with voicing. Several proposed acoustic correlates to perceptual features of stop consonant-vowel syllables, including the initial spectrum, formant transitions, and voice-onset time, are represented in the temporal properties of auditory-nerve fiber responses. Nonlinear suppression affects the temporal features of the responses, particularly those of low/medium-spontaneous-rate fibers.

Acoustic Stimulation

Frequency selectivity of single cochlear-nerve fibers based on the temporal response pattern to two-tone signals.

The physiological basis of auditory frequency selectivity was investigated by recording the temporal response patterns of single cochlear-nerve fibers in the cat. The characteristic frequency and sharpness of tuning was determined for low-frequency cochlear-nerve fibers with two-tone signals whose frequency components were of equal amplitude and starting phase. The measures were compared with those obtained with sinusoidal signals. The two-tone characteristic frequency (2TCF) is defined as the arithmetic-center frequency at which the fiber is synchronized to both signal frequencies in equal measure. The 2TCF closely corresponds to the characteristic frequency as determined by the frequency threshold curve. Moreover, the 2TCF changes relatively little (2%-12%) over a 60-dB intensity range. The 2TCF generally shifts upward with increasing intensity for cochlear-nerve fibers tuned to frequencies below 1 kHz and shifts downward as a function of intensity for units with characteristic frequencies (CF's) above 1 kHz. The shifts in the 2TCF are considerably smaller than those observed with sinusoidal signals. Filter functions were derived from the synchronization pattern to the two-tone signal by varying the frequency of one of the components over the fiber's response area while maintaining the other component at the 2TCF. The frequency selectivity of the two-tone filter function was determined by dividing the vector strength to the variable frequency signal by the vector strength to the CF tone. The filter function was measured 10 dB down from the peak (2T Q 10 dB) and compared with the Q 10 dB of the frequency threshold curve. The correlation between the two measures of frequency selectivity was 0.72. The 2T Q 10 dB does change as a function of intensity. The magnitude and direction of the change is dependent on the sharpness of tuning at low and moderate sound-pressure levels (SPL's). The selectivity of the more sharply tuned fibers (2T Q 10 dB greater than 3) diminishes at intensities above 60 dB SPL. However, the broadening of selectivity is relatively small in comparison to discharge rate-based measures of selectivity. The selectivity of the more broadly tuned units remains unchanged or improves slightly at similar intensity levels. The present data indicate that the frequency selectivity and tuning of low-frequency cochlear-nerve fibers are relatively stable over a 60-dB range of SPL's when measured in terms of their temporal discharge properties.

Animals

A two-stage nonlinear cochlear model possesses automatic gain control.

A model of the cochlea is explored using as stimuli two simultaneously presented sinusoids of equal amplitude. The model consists of two stages: a linear bandpass filter, followed by a reservoir-type representation of the hair-cell/nerve-fiber complex. Fast Fourier transforms of the model's output were computed. While the amplitudes of the individual response components were strongly nonlinear functions of intensity, the ratio of the magnitudes of the response components at the frequencies of the two stimulating sinusoids was found to be nearly equal, over a wide intensity range, to the ratio of the amplitudes which those stimulating sinusoids possessed at the output of the filter. Thus the reservoir stage exerts "automatic gain control".

Adaptation, Physiological

Thresholds for primary auditory fibers using statistically defined criteria.

The discharge behavior of auditory-nerve fibers near "threshold" was investigated in anesthetized cats using low-intensity sinusoidal stimuli presented at the respective characteristic frequencies. Particular attention was paid to fibers with "low" and "medium" rates. Estimates of threshold derived from statistically significant increases in discharge rate indicate that the average threshold values for low-spontaneous fibers are only slightly higher (ca. 5 dB) than the averages for the corresponding high-spontaneous fibers, with the medium-spontaneous fibers having intermediate averages. The difference between these average values is considerably less than the more than 20-dB difference obtained using threshold criteria based on an absolute increment in discharge rate [e.g., M. C. Liberman, J. Acoust. Soc. Am. 63, 442-455 (1978)]. The main reason for the difference between the results of the two techniques is the fact that the slopes of the rate-intensity functions for the high-spontaneous fibers are considerably steeper near "threshold" than those for fibers of the other two classes. The results are taken as supportive of a recent model of primary-fiber discharge [C. D. Geisler, Brain Res. 212, 198-201 (1981)].

Animals