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Marcel van der Heijden

Publications and source records attributed to Marcel van der Heijden.

9 recordsLinked to original sources

Correlation index: a new metric to quantify temporal coding.

The standard procedure to study temporal encoding of sound waveforms in the auditory system has been Fourier analysis of responses to periodic stimuli. We introduce a new metric--correlation index (CI)--which is based on a simple counting of spike coincidences. It can be used for responses to aperiodic stimuli and does not require knowledge of the stimulus. Moreover, the basic procedure of comparing spiketimes in spiketrains is more physiological than currently used methods for temporal analysis. The CI is the peak value of the normalized shuffled autocorrelogram (SAC), which provides a quantitative summary of temporal structure in the neural response to arbitrary stimuli. We illustrate the CI and SACs by comparing temporal coding in the auditory nerve and output fibers of the cochlear nucleus.

Acoustic Stimulation↗

Auditory midbrain and nerve responses to sinusoidal variations in interaural correlation.

The human sensitivity to interaural temporal differences in the acoustic waveforms to the two ears shows remarkable acuity but is also very sluggish. Fast changes in binaural parameters are not detectable, and this inability contrasts sharply with the excellent temporal resolution of the monaural auditory system. We studied the response of binaural neurons in the inferior colliculus of the cat to sinusoidal changes in the interaural correlation of broadband noise. Responses to the same waveforms were also obtained from auditory nerve fibers and further analyzed with a coincidence analysis. Overall, the auditory nerve and inferior colliculus showed a similar ability to code changes in interaural correlation. This ability extended to modulation frequencies an order of magnitude higher than the highest frequencies detected binaurally in humans. We conclude that binaural sluggishness is not caused by a lack of temporal encoding of fast binaural changes at the level of the midbrain. We hypothesize that there is no neural substrate at the level of the midbrain or higher to read out this temporal code and that this constitutes a low-pass "sluggishness" filter.

Acoustic Stimulation↗

Decorrelation sensitivity of auditory nerve and anteroventral cochlear nucleus fibers to broadband and narrowband noise.

Binaural neurons show remarkable sensitivity to temporal differences in the waveforms at the two ears. This ability obviously requires temporal coding of sound waveforms in the monaural afferents that converge on such binaural neurons. We introduce a new analysis to investigate how well responses of single monaural neurons support discrimination of decorrelations in waveforms. Spike trains from auditory nerve (AN) and anteroventral cochlear nucleus (AVCN) neurons of cats to many repetitions of a set of broadband and narrowband noise tokens were obtained. The normalized correlation between the noise tokens ranged from 0.99 to -1. A coincidence and signal detection analysis was used to perform a correlation discrimination task using the monaural spike trains. The correlation discrimination thresholds derived from AVCN neurons were lower than those derived from AN fibers and sometimes as low as human psychophysical just noticeable differences. Importantly, low detection thresholds required comparisons of spike trains at small internal delays. Bandwidth dependence of neural decorrelation thresholds agreed with psychophysical data when large internal delays contributed to the detection. We conclude that, in the context of correlation discrimination, coding by AVCN fibers is superior to that by AN fibers and that these discriminations require a distribution of internal or best delays in binaural processing that differs from the predictions from studies of discrimination in interaural time delays.

Acoustic Stimulation↗

Enhanced temporal response properties of anteroventral cochlear nucleus neurons to broadband noise.

Compared with auditory nerve (AN) fibers, trapezoid body (TB) fibers of the cat show enhanced synchronization to low-frequency tones. This phenomenon probably contributes to the high temporal resolution of binaural processing. We examined whether enhanced synchronization also occurs to sustained broadband noise. We recorded responses to a reference Gaussian noise and its polarity-inverted version in the TB of barbiturate-anesthetized cats. From these we constructed shuffled autocorrelograms (SACs) and quantified spike timing by measuring the amplitude and width of their central peak. Many TB fibers with low characteristic frequency (CF) showed SACs with higher and narrower central peaks than ever observed in the AN, indicating better consistency and precision of temporal coding. Larger peaks were also observed in TB fibers with high CF, but this was mostly caused by higher average firing rates, resulting in a larger number of coincident spikes across stimulus repetitions. The results document monaural preprocessing of the temporal information delivered to binaural nuclei in the olivary complex, which likely contributes to the high sensitivity to interaural time differences.

Acoustic Stimulation↗

Cochlear gain control.

The nonlinear auditory phenomena of compression, suppression, and distortion are known to have a cochlear-mechanical origin. An instantaneous nonlinear transfer function is often assumed to underlie these phenomena, but there are experimental indications that auditory nonlinearity is sluggish rather than instantaneous. This study analyzes the consequences of such sluggishness, using automatic gain control (AGC) as a model noninstantaneous nonlinearity. The distinctive characteristic of AGC, its delayed action, is shown to produce a number of observable and measurable effects that distinguish AGC from instantaneous nonlinearities. A major class of such AGC-specific effects concerns the phase of aural distortion products. For example, the phase of the cancellation tone in the classical psychoacoustic cancellation paradigm is linearly related to the frequency spacing of the primary tones in an AGC, as opposed to the square-law relationship produced by an instantaneous nonlinearity. These and other predictions are confronted with experimental data from the literature. The impact of putative AGC-related delays on the interpretation of distortion product otoacoustic emissions (DPOAEs) is discussed. Detailed suggestions are made for experiments specifically aimed at determining whether cochlear nonlinearity is instantaneous or delayed.

Animals↗

Temporal damping in response to broadband noise. I. Inferior colliculus.

Many cells in the inferior colliculus (IC) are sensitive to interaural time differences (ITDs), in the form of an oscillatory dependency of average firing rate on ITD. We studied the degree of damping in such binaural responses, recording from neurons in the inferior colliculus of pentobarbital-anesthetized cats to binaural broadband noise and tones. Noise-delay functions and composite curves were characterized by computing the difference between responses to correlated and anticorrelated stimuli. We use a new metric, based on the envelope of this difference, to quantify damping. There is a clear relationship between damping and characteristic frequency (CF), but even neurons of the same CF can differ in their damping. For individual cells, damping can be stronger to tones or to noise; at the population level the two are positively correlated and are scarcely affected by SPL. The frequencies that dominate ITD sensitivity are near the CF in response to noise, but are often below CF in response to tones. These findings qualify conclusions from earlier reports but overall they support the conclusion that, at a population level, basic aspects of binaural responses to wideband noise are consistent with summed responses to pure tones.

Acoustic Stimulation↗

The speed of auditory low-side suppression.

The nonlinear cochlear phenomenon of two-tone suppression is known to be very fast, but precisely how fast is unknown. We studied the timing of low-side suppression in the auditory nerve of the cat using multitone complexes as auditory stimuli. An evalution of the group delays of the responses to these complexes allowed us to measure the timing of the responses with sub-millisecond accuracy for a large number of fibers with characteristic frequencies (CFs) between 2 and 40 kHz. In particular, we measured the delays with which the same below-CF tone complexes affected the response either as an excitor (when presented alone) or as a suppressor (when combined with a CF probe). For CFs <10 kHz, we found that the delay of suppression was larger than the delay of excitation by several hundred microseconds. The difference between the delay of suppression and that of excitation decreased with increasing CF, becoming negligible for CFs >15 kHz. The results are analyzed in terms of traveling-wave delays and a purported cochlear gain control. The data suggest that suppression originates from a gain-control mechanism with an integration time in the order of two cycles of CF.

Acoustic Stimulation↗

Temporal properties of responses to broadband noise in the auditory nerve.

Temporal information in the responses of auditory neurons to sustained sounds has been studied mostly with periodic stimuli, using measures that are based on Fourier analysis. Less information is available on temporal aspects of responses to nonperiodic wideband sounds. We recorded responses to a reference Gaussian noise and its polarity-inverted version in the auditory nerve of barbiturate-anesthetized cats and used shuffled autocorrelograms (SACs) to quantify spike timing. Two metrics were extracted from the central peak of autocorrelograms: the peak-height and the width at halfheight. Temporal information related to stimulus fine-structure was isolated from that to envelope by subtracting or adding responses to the reference and inverted noise. Peak-height and halfwidth generally behaved as expected from the existing body of data on phase-locking to pure tones and sinusoidally amplitude-modulated tones but showed some surprises as well. Compared with synchronization to low-frequency tones, SACs reveal large differences in temporal behavior between the different classes of nerve fibers (based on spontaneous rate) as well as a strong dependence on characteristic frequency (CF) throughout the phase-locking range. SACs also reveal a larger temporal consistency (i.e., tendency to discharge at the same point in time on repeated presentation of the same stimulus) in the responses to the stochastic noise stimulus than in the responses to periodic tones. Responses at high CFs reflect envelope phase-locking and are consistent with previous reports using sinusoidal AM. We conclude that the combined use of broadband noise and SAC analysis allow a more general characterization of temporal behavior than periodic stimuli and Fourier analysis.

Acoustic Stimulation↗

Cochlear phase and amplitude retrieved from the auditory nerve at arbitrary frequencies.

It is currently impossible to mechanically measure the overall vibration pattern of the intact mammalian cochlea because of its inaccessibility and vulnerability. At first sight, data from the auditory nerve are a poor substitute because of their limited temporal resolution. The nonlinear character of neural coding, however, causes low-frequency interactions among the components of multitone stimuli. We designed a novel stimulus for which these interactions take a particularly systematic form, and we recorded the response of the auditory nerve to this stimulus. A careful analysis of interactions in the data allowed us to reconstruct frequency transfer functions (both their amplitude and their phase) at multiple points spanning the entire length of the cochlea. The generic character of our stimuli and analysis suggests its wider use in nonlinear system analysis, particularly in those instances in which limitations in temporal resolution restrict the use of customary methods.

Acoustic Stimulation↗