Biomedical subjects

Jesko L Verhey

Publications and source records attributed to Jesko L Verhey.

11 recordsLinked to original sources

PubMed2006

The temporal representation of the delay of iterated rippled noise with positive or negative gain by chopper units in the cochlear nucleus.

The role of chopper units in representing the pitch of complex sounds is unresolved. Traditionally chopper units have been regarded as primarily responding to the stimulus envelope of complex stimuli. This has been supported by the response of chopper units to iterated rippled noise (IRN) as they can provide a robust representation of the delay of IRN with positive gain (+) in their first-order interspike intervals and for some chopper units this representation is relatively level independent. The envelope modulation of IRN(+), and pitch, is at the reciprocal of the delay, the pitch of IRN with negative gain (IRN(-)) is often at twice the delay. This distinction between IRN(+) and IRN(-) can be used to help determine whether a unit is simply responding to modulation or to stimulus fine structure. Chopper units with relatively high best frequencies (BF) are unable to represent the distinction between IRN(+) and IRN(-). However, in this study it is shown that at least some chopper units, with low BFs (<1.25 kHz), can represent the pitch of the IRN(-) as perceived perceptually.

Animals↗

PubMed2006

Spectral loudness summation as a function of duration for hearing-impaired listeners.

Spectral loudness summation was measured for 10- and 1000-ms long bandpass-noise signals with bandwidths in the range of 0.2 to 6.4 kHz centered at 2 kHz for nine listeners with hearing impairment of primarily cochlear origin. Loudness matches between signals with the same duration and different bandwidth were obtained using an adaptive two-interval, two-alternative forced-choice procedure with interleaved tracks. The 3200-Hz wide reference signal had a level of 45 dB or 65 dB SPL. Every signal was individually frequency equalized prior to presentation in order to ensure audibility of all spectral components for the two reference input levels. Generally the same amount of spectral loudness summation for 10- and 1000-ms long signals was obtained for both reference levels. However, some hearing-impaired listeners show, similarly to normal-hearing listeners, a larger spectral loudness summation for short than for long duration signals, indicating that duration effects in spectral loudness summation are caused by retrocochlear processes.

Acoustic Stimulation↗

PubMed2006

Spectral loudness summation for short and long signals as a function of level.

The equal-loudness-ratio hypothesis (ELRH) assumes that the loudness ratio between equal-level long and short signals with the same spectrum is independent of level and spectrum. Thus, it predicts the same amount of spectral loudness summation for equal-level short and long signals. To investigate how duration and level affect spectral loudness summation, loudness functions for 10- and 1000-ms bandpass-filtered noise centered at 2 kHz were obtained for normal-hearing listeners using categorical loudness scaling. The bandwidth was varied from 200 to 6400 Hz. From the loudness functions the level difference between the 3200-Hz-wide reference signal and the equally loud test signals was determined for reference levels in the range from 25 to 105 dB SPL. In addition, for two reference levels (45 and 65 dB SPL), this level difference was measured using a loudness-matching procedure. Both procedures showed similar results for the two reference levels. Over a wide level range spectral loudness summation is larger for the short than for the equal-level long signals. This is in contrast to the predictions of the ELRH. However, the data at moderate levels are consistent with the expectations of a modified ELRH which assumes a smaller loudness ratio for broadband than for narrow-band signals.

Adult↗

PubMed2006

Role of suppression and retro-cochlear processes in comodulation masking release.

Recent physiological studies suggest that comodulation masking release (CMR) could be a consequence of wideband inhibition at the level of the cochlear nucleus. The present study investigates whether the existence region of psychophysical CMR is comparable to the inhibitory areas of units showing a physiological correlate of CMR. Since the inhibitory areas are similar to suppressive regions at the level of the basilar membrane, the amount of CMR that can be accounted for by suppression was determined by predicting the data with a model incorporating a peripheral nonlinearity. A CMR of up to 6 dB could still be experimentally observed for a flanking band (FB) four octaves below the on-frequency masker (OFM). For FB frequencies below the OFM, the suggested model predicts CMR equal to the measured CMR for high levels of the FB. The model underestimates the magnitude of CMR for midlevels of the FB, indicating that suppression alone cannot account for CMR. The data are consistent with the hypothesis that wideband inhibition plays a role in CMR.

Adult↗

PubMed2005

Temporal representation of the delay of iterated rippled noise in the dorsal cochlear nucleus.

It has been suggested that the dorsal cochlear nucleus (DCN) is involved in the temporal representation of both envelope periodicity and pitch. This hypothesis is tested using iterated rippled noise (IRN), which is generated by a cascade of delay and add [IRN(+)] or delay and subtract [IRN(-)] operations. The autocorrelation functions (ACFs) of the waveform and the envelope of IRN(+) have a first peak at the delay, which corresponds to the perceived pitch of the IRN. With the same delay, the pitch of IRN(-) is generally an octave lower than for IRN(+). This is reflected in a first peak at twice the delay in the ACF of the waveform for IRN(-). In contrast, for identical delays, the ACF of the envelope for both IRN(-) and IRN(+) is the same. Thus the use of IRN allows the distinction between envelope- or fine-structure sensitivity. Recordings were made from 135 single units (BFs <5 kHz) in the DCN of the anesthetized guinea pig using IRN with delays ranging from 1 to 32 ms. In our sample 42% were sensitive to the periodicity of IRN(+) and were tuned to a particular delay in their first-order interspike interval histograms (ISIHs). This tuning was highly correlated with their response to white noise. Most units with best frequencies (BFs) <500 Hz show a different all-order ISIH for IRN(+) and IRN(-), which corresponds to the perceived pitch difference, whereas units with higher BFs show a similar response to IRN(+) and IRN(-). The results indicate that low-frequency units (BF <500 Hz) in the DCN may be involved in the representation of the waveform fine structure, although units with BFs >500 Hz are able to encode only the envelope periodicity of broadband IRN in their temporal discharge characteristics.

Acoustic Stimulation↗

PubMed2004

Responses of dorsal cochlear nucleus neurons to signals in the presence of modulated maskers.

The detection of a signal in noise is enhanced when the masking noise is coherently modulated over a wide range of frequencies. This phenomenon, known as comodulation masking release (CMR), has been attributed to across-channel processing; however, the relative contribution of different stages in the auditory system to such across-channel processing is unknown. It has been hypothesized that wideband or lateral inhibition may underlie a physiological correlate of CMR. To further test this hypothesis, we have measured the responses of single units from the dorsal cochlear nucleus in which wideband inhibition is particularly pronounced. Using a sinusoidally amplitude-modulated tone at the best frequency of each unit as a masker, a pure-tone signal was added in the dips of the masker modulation. Flanking bands (FBs, also amplitude-modulated pure tones) were positioned to fall within the inhibitory sidebands of the receptive field of the unit. The FBs were either in phase (comodulated) or out of phase (codeviant) with the on-frequency masker. For the majority of units, the addition of the comodulated FBs produced a strong reduction in the response to the masker modulation, making the signal more salient in the post stimulus time histograms. The change in spike rate in response to the signal between the masker and signal-plus-masker conditions was greatest for the comodulated condition for 29 of 45 units. These results are consistent with the hypothesis that wideband inhibition may play a role in across-channel processing at an early stage in the auditory pathway.

Acoustic Stimulation↗

PubMed2003

The psychophysics and physiology of comodulation masking release.

The ability to detect auditory signals from background noise may be enhanced by the addition of energy in frequency regions well removed from the frequency of the signal. However, it is important that this energy is amplitude-modulated in a coherent way across frequencies, i.e. comodulated. This enhancement of signal detectability is known as comodulation masking release (CMR), and in this review we show that CMR is largest if: (1) the total masker's bandwidth is large, (2) the modulation frequency is low, (3) the modulation depth is high, (4) the envelope is regular and, (5) the masker's spectrum level is high. Possible physiological correlates of CMR have been found at different levels of the auditory pathway. Current hypotheses for the underlying physiological mechanisms, including wide-band inhibition or the disruption of masker modulation envelope response, are discussed.

Action Potentials↗

PubMed2003

Modulation masking produced by complex tone modulators.

Thresholds were measured for detecting sinusoidal amplitude modulation in the presence of a complex-tone masker modulation. Both modulations were applied to the same sinusoidal carrier. Two different masker modulations were used: (i) a pair of components beating at the difference frequency and (ii) a three-tone complex producing a sinusoidal amplitude modulation of the modulation depth at the difference frequency between adjacent components. Both maskers show a periodicity in the waveform that is not contained in the envelope spectrum itself but can be observed when the envelope of the envelope, referred to as the "venelope" [Ewert et al., J. Acoust. Soc. Am. 112, 2921-2931 (2002)], is calculated. For a signal frequency equal to the masker-venelope periodicity, modulation depth at threshold was measured as a function of the signal phase relative to the phase of the masker-venelope component. Signal frequencies of 5, 30, and 90 Hz were used. It was found that masking was phase dependent for all three signal frequencies. Thresholds were lower for the in-phase condition, where maxima in the signal waveform coincided with maxima in the masker-venelope waveform, than for the antiphase condition. The maximum threshold difference was 15 dB. The results are in contrast to recent data [Moore et al., J. Acoust. Soc. Am. 106, 908-918 (1999)], where lowest thresholds were reported for the antiphase condition in a similar experiment. The present data are in line with the idea that a nonlinearity prior to a modulation filterbank extracts the venelope of the masker modulator. However, a compressive nonlinearity such as that associated with the processing on the basilar membrane cannot account for the empirical findings, since it predicts the opposite phase effect.

Adult↗

PubMed2002

Modeling the influence of inherent envelope fluctuations in simultaneous masking experiments.

Masked thresholds are measured and simulated for bandpass-noise signals ranging in bandwidth from 4 to 256 Hz in the presence of a masking bandpass noise also ranging in bandwidth from 4 to 256 Hz. Signal and masker are centered at 2 kHz. To investigate the role of temporal processing in simultaneous masking, simulations were performed with the modulation-filterbank model by Dau et al. [J. Acoust. Soc Am. 102, 2906-2919 (1997)]. For a fixed masker bandwidth, thresholds are independent of the signal bandwidth as long as the signal bandwidth does not exceed the masker bandwidth and thresholds decrease with increasing masker bandwidth in those conditions. A simple modulation-low-pass filter (energy integrator) would be sufficient to describe the experimental results in those conditions. In contrast, the processing by a modulation filterbank is necessary to account for the conditions of "asymmetry of masking," where thresholds for signals with bandwidths larger than the masker bandwidth are much lower than those for the reversed condition. In those conditions, the modulation-filterbank model is able to use the inherent higher modulation frequencies of the signal as an additional cue.

Adult↗

PubMed2002

Spectral loudness summation as a function of duration.

Loudness was measured as a function of signal bandwidth for 10-, 100-, and 1000-ms-long signals. The test and reference signals were bandpass-filtered noise spectrally centered at 2 kHz. The bandwidth of the test signal was varied from 200 to 6400 Hz. The reference signal had a bandwidth of 3200 Hz. The reference levels were 45, 55, and 65 dB SPL. The level to produce equal loudness was measured with an adaptive, two-interval, two-alternative forced-choice procedure. A loudness matching procedure was used, where the tracks for all signal pairs to be compared were interleaved. Mean results for nine normal-hearing subjects showed that the magnitude of spectral loudness summation depends on signal duration. For all reference levels, a 6- to 8-dB larger level difference between equally loud signals with the smallest (delta f = 200 Hz) and largest (delta f = 6400 Hz) bandwidth is found for 10-ms-long signals than for the 1000-ms-long signals. The duration effect slightly decreases with increasing reference loudness. As a consequence, loudness models should include a duration-dependent compression stage. Alternatively, if a fixed loudness ratio between signals of different duration is assumed, this loudness ratio should depend on the signal spectrum.

Adult↗

PubMed2002

Spectro-temporal processing in the envelope-frequency domain.

The frequency selectivity for amplitude modulation applied to tonal carriers and the role of beats between modulators in modulation masking were studied. Beats between the masker and signal modulation as well as intrinsic envelope fluctuations of narrow-band-noise modulators are characterized by fluctuations in the "second-order" envelope (referred to as the "venelope" in the following). In experiment 1, masked threshold patterns (MTPs), representing signal modulation threshold as a function of masker-modulation frequency, were obtained for signal-modulation frequencies of 4, 16, and 64 Hz in the presence of a narrow-band-noise masker modulation, both applied to the same sinusoidal carrier. Carrier frequencies of 1.4, 2.8, and 5.5 kHz were used. The shape and relative bandwidth of the MTPs were found to be independent of the signal-modulation frequency and the carrier frequency. Experiment 2 investigated the extent to which the detection of beats between signal and masker modulation is involved in tone-in-noise (TN), noise-in-tone (NT), and tone-in-tone (TT) modulation masking, whereby the TN condition was similar to the one used in the first experiment. A signal-modulation frequency of 64 Hz, applied to a 2.8-kHz carrier, was tested. Thresholds in the NT condition were always lower than in the TN condition, analogous to the masking effects known from corresponding experiments in the audio-frequency domain. TT masking conditions generally produced the lowest thresholds and were strongly influenced by the detection of beats between the signal and the masker modulation. In experiment 3, TT masked-threshold patterns were obtained in the presence of an additional sinusoidal masker at the beat frequency. Signal-modulation frequencies of 32, 64, and 128 Hz, applied to a 2.8-kHz carrier, were used. It was found that the presence of an additional modulation at the beat frequency hampered the subject's ability to detect the envelope beats and raised thresholds up to a level comparable to that found in the TN condition. The results of the current study suggest that (i) venelope fluctuations play a similar role in modulation masking as envelope fluctuations do in spectral masking, and (ii) envelope and venelope fluctuations are processed by a common mechanism. To interpret the empirical findings, a general model structure for the processing of envelope and venelope fluctuations is proposed.

Adult↗