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Christophe Micheyl

Publications and source records attributed to Christophe Micheyl.

At least 19 recordsLinked to original sources

Cortical FMRI activation to sequences of tones alternating in frequency: relationship to perceived rate and streaming.

Human listeners were functionally imaged while reporting their perception of sequences of alternating-frequency tone bursts separated by 0, 1/8, 1, or 20 semitones. Our goal was to determine whether functional magnetic resonance imaging (fMRI) activation of auditory cortex changes with frequency separation in a manner predictable from the perceived rate of the stimulus. At the null and small separations, the tones were generally heard as a single stream with a perceived rate equal to the physical tone presentation rate. fMRI activation in auditory cortex was appreciably phasic, showing prominent peaks at the sequence onset and offset. At larger-frequency separations, the higher- and lower-frequency tones perceptually separated into two streams, each with a rate equal to half the overall tone presentation rate. Under those conditions, fMRI activation in auditory cortex was more sustained throughout the sequence duration and was larger in magnitude and extent. Phasic to sustained changes in fMRI activation with changes in frequency separation and perceived rate are comparable to, and consistent with, those produced by changes in the physical rate of a sequence and are far greater than the effects produced by changing other physical stimulus variables, such as sound level or bandwidth. We suggest that the neural activity underlying the changes in fMRI activation with frequency separation contribute to the coding of the co-occurring changes in perceived rate and perceptual organization of the sound sequences into auditory streams.

Acoustic Stimulation↗

Influence of musical and psychoacoustical training on pitch discrimination.

This study compared the influence of musical and psychoacoustical training on auditory pitch discrimination abilities. In a first experiment, pitch discrimination thresholds for pure and complex tones were measured in 30 classical musicians and 30 non-musicians, none of whom had prior psychoacoustical training. The non-musicians' mean thresholds were more than six times larger than those of the classical musicians initially, and still about four times larger after 2h of training using an adaptive two-interval forced-choice procedure; this difference is two to three times larger than suggested by previous studies. The musicians' thresholds were close to those measured in earlier psychoacoustical studies using highly trained listeners, and showed little improvement with training; this suggests that classical musical training can lead to optimal or nearly optimal pitch discrimination performance. A second experiment was performed to determine how much additional training was required for the non-musicians to obtain thresholds as low as those of the classical musicians from experiment 1. Eight new non-musicians with no prior training practiced the frequency discrimination task for a total of 14 h. It took between 4 and 8h of training for their thresholds to become as small as those measured in the classical musicians from experiment 1. These findings supplement and qualify earlier data in the literature regarding the respective influence of musical and psychoacoustical training on pitch discrimination performance.

Acoustic Stimulation↗

Detection and F0 discrimination of harmonic complex tones in the presence of competing tones or noise.

Normal-hearing listeners' ability to "hear out" the pitch of a target harmonic complex tone (HCT) was tested with simultaneous HCT or noise maskers, all bandpass-filtered into the same spectral region (1200-3600 Hz). Target-to-masker ratios (TMRs) necessary to discriminate fixed fundamental-frequency (F0) differences were measured for target F0s between 100 and 400 Hz. At high F0s (400 Hz), asynchronous gating of masker and signal, presenting the masker in a different F0 range, and reducing the F0 rove of the masker, all resulted in improved performance. At the low F0s (100 Hz), none of these manipulations improved performance significantly. The findings are generally consistent with the idea that the ability to segregate sounds based on cues such as F0 differences and onset/offset asynchronies can be strongly limited by peripheral harmonic resolvability. However, some cases were observed where perceptual segregation appeared possible, even when no peripherally resolved harmonics were present in the mixture of target and masker. A final experiment, comparing TMRs necessary for detection and F0 discrimination, showed that F0 discrimination of the target was possible with noise maskers at only a few decibels above detection threshold, whereas similar performance with HCT maskers was only possible 15-25 dB above detection threshold.

Acoustic Stimulation↗

Likelihood ratio, optimal decision rules, and correct response probabilities in a signal detection theoretic, equal-variance Gaussian model of the observer in the 4IAX paradigm.

This article provides a synthetic account of the likelihood ratio, optimal decision rules, and correct response probabilities in a signal detection theoretic model of the observer in the dual-pair comparison, or four-interval AX (4IAX), paradigm. The model assumes a static sampling process, resulting in four, equal-variance normally distributed (i.e., Gaussian) observations on each trial First, a likelihood ratio equation allowing for an arbitrary degree of correlation between observations is provided. Specific solutions for the cases of independent and highly correlated observations are then derived. It is shown that these solutions, and the associated decision rules, correspond to those provided independently in earlier publications. A modified 4IAX paradigm involving, as a standard, an additional stimulus (C) located medially between the A and the B stimuli is also considered. It is shown that the optimal (static, equal-variance, Gaussian) decision model for this paradigm is unaffected by correlation between observations and is equivalent to the standard 4IAX with highly correlated observations. Finally, we consider how, under the considered (static, equal-variance, Gaussian) model, the proportion of correct responses in the different versions of the 4IAX paradigm is related to d', and a solution for the case of independent observations is provided.

Decision Making↗

Perceptual organization of tone sequences in the auditory cortex of awake macaques.

Acoustic sequences such as speech and music are generally perceived as coherent auditory "streams," which can be individually attended to and followed over time. Although the psychophysical stimulus parameters governing this "auditory streaming" are well established, the brain mechanisms underlying the formation of auditory streams remain largely unknown. In particular, an essential feature of the phenomenon, which corresponds to the fact that the segregation of sounds into streams typically takes several seconds to build up, remains unexplained. Here, we show that this and other major features of auditory-stream formation measured in humans using alternating-tone sequences can be quantitatively accounted for based on single-unit responses recorded in the primary auditory cortex (A1) of awake rhesus monkeys listening to the same sound sequences.

Acoustic Stimulation↗

Generalization of frequency discrimination learning across frequencies and ears: implications for underlying neural mechanisms in humans.

Frequency discrimination thresholds (FDTs) at 750, 1500, 3000, and 6000 Hz were measured in 32 normal-hearing listeners before and after each listener practiced the task for 12 h at one of the above frequencies using a single ear. Marked improvements in thresholds taking place over several hours were observed during the frequency- and ear-specific training period. Comparisons between pre- and posttraining thresholds showed large improvements at the trained frequency, but also at other frequencies. The improvements were initially slightly-but significantly-larger at the trained frequency than at untrained frequencies. However, this trained-frequency advantage disappeared rapidly during the course of the two-hour multifrequency posttraining session, suggesting rapid relearning or learning generalization across frequencies. In contrast, no significant ear specificity was found, not even at early stages of the posttraining session. These findings add to earlier results suggesting that, in humans, frequency discrimination learning is only weakly frequency-specific, and they reveal that a complete generalization across frequencies can occur rapidly with little retraining at the initially untrained frequencies. Implications regarding underlying mechanisms are discussed.

Adult↗

Neuromagnetic correlates of streaming in human auditory cortex.

The brain is constantly faced with the challenge of organizing acoustic input from multiple sound sources into meaningful auditory objects or perceptual streams. The present study examines the neural bases of auditory stream formation using neuromagnetic and behavioral measures. The stimuli were sequences of alternating pure tones, which can be perceived as either one or two streams. In the first experiment, physical stimulus parameters were varied between values that promoted the perceptual grouping of the tone sequence into one coherent stream and values that promoted its segregation into two streams. In the second experiment, an ambiguous tone sequence produced a bistable percept that switched spontaneously between one- and two-stream percepts. The first experiment demonstrated a strong correlation between listeners' perception and long-latency (>60 ms) activity that likely arises in nonprimary auditory cortex. The second demonstrated a covariation between this activity and listeners' perception in the absence of physical stimulus changes. Overall, the results indicate a tight coupling between auditory cortical activity and streaming perception, suggesting that an explicit representation of auditory streams may be maintained within nonprimary auditory areas.

Acoustic Stimulation↗

Comparing F0 discrimination in sequential and simultaneous conditions.

In an influential study, Carlyon and Shackleton [J. Acoust. Soc. Am. 95, 3541-3554 (1994)] measured listeners' performance (d') in fundamental-frequency (F0) discrimination between harmonic complex tones (HCTs) presented simultaneously in different spectral regions and compared their performance with that found in a sequential-comparison task. In this Letter, it is suggested that Carlyon and Shackleton's analysis of the simultaneous-comparison data did not adequately reflect their assumption that listeners were effectively comparing F0's across regions. A reanalysis consistent with this assumption is described. The new results suggest that under the assumption that listeners were effectively comparing F0 across regions, their performance in this task was substantially higher than originally estimated by Carlyon and Shackleton, and in some conditions much higher than expected from the performances measured in a traditional F0-discrimination task with sequential HCTs. Possible explanations for this outcome, as well as alternative interpretations, are proposed.

Discrimination, Psychological↗

Sequential F0 comparisons between resolved and unresolved harmonics: no evidence for translation noise between two pitch mechanisms.

Carlyon and Shackleton [J. Acoust. Soc. Am. 95, 3541-3554 (1994)] suggested that fundamental-frequency (F0) discrimination performance between resolved and unresolved harmonics is limited by an internal "translation" noise between the outputs of two distinct F0 encoding mechanisms, in addition to the encoding noise associated with each mechanism. To test this hypothesis further, F0 difference limens (DLF0s) were measured in six normal-hearing listeners using sequentially presented groups of harmonics. The two groups of harmonics presented on each trial were bandpass filtered into the same or different spectral regions, in such a way that both groups contained mainly resolved harmonics, both groups contained only unresolved harmonics, or one group contained mainly resolved and the other only unresolved harmonics. Three spectral regions (low: 600-1150 Hz, mid: 1400-2500 Hz, or high: 3000-5250 Hz) and two nominal F0s (100 and 200 Hz) were used. The DLF0s measured in across-region conditions were well accounted for by a model assuming only two sources of internal noise: the encoding noise estimated on the basis of the within-region results plus a constant noise associated with F0 comparisons across different spectral regions, independent of resolvability. No evidence for an across-pitch-mechanism translation noise was found. A reexamination of previous evidence for the existence of such noise suggests that the present negative outcome is unlikely to be explained by insufficient measurement sensitivity or an unusually large across-region comparison noise in the present study. While the results do not rule out the possibility of two separate pitch mechanisms, they indicate that the F0s of sequentially presented resolved and unresolved harmonics can be compared internally at no or negligible extra cost.

Acoustic Stimulation↗

Auditory processing of real and illusory changes in frequency modulation (FM) phase.

Auditory processing of frequency modulation (FM) was explored. In experiment 1, detection of a tau-radians modulator phase shift deteriorated as modulation rate increased from 2.5 to 20 Hz, for 1- and 6-kHz carriers. In experiment 2, listeners discriminated between two 1-kHz carriers, where, mid-way through, the 10-Hz frequency modulator had either a phase shift or increased in depth by deltaD% for half a modulator period. Discrimination was poorer for deltaD = 4% than for smaller or larger increases. These results are consistent with instantaneous frequency being smoothed by a time window with a total duration of about 110 ms. In experiment 3, the central 200-ms of a 1-s 1-kHz carrier modulated at 5 Hz was replaced by noise, or by a faster FM applied to a more intense 1-kHz carrier. Listeners heard the 5-Hz FM continue at the same depth throughout the stimulus. Experiments 4 and 5 showed that, after an FM tone had been interrupted by a 200-ms noise, listeners were insensitive to the phase at which the FM resumed. It is argued that the auditory system explicitly encodes the presence, and possibly the rate and depth, of FM in a way that does not preserve information on FM phase.

Auditory Threshold↗

Enhanced frequency discrimination near the hearing loss cut-off: a consequence of central auditory plasticity induced by cochlear damage?

Patients with steeply sloping hearing losses of cochlear origin may exhibit enhanced difference limens for frequency (DLFs) near the cut-off frequency (Fc) of their hearing loss. This effect has been related to observations in deafened animals of an over-representation of Fc in the primary auditory cortex. However, alternative interpretations in terms of peripheral mechanisms have not been eliminated. In the present study, we assessed the possible role of two peripheral mechanisms [loudness cues and spontaneous otoacoustic emissions (SOAEs)] in a group of patients with high-frequency hearing loss. We tested first whether the DLF enhancement effect was still observed under conditions where subjects could not rely on loudness cues to perform the frequency discrimination task. To achieve this, we adjusted the nominal level of each stimulus so that it fell on an equal loudness contour measured at very fine (1/8 octave) frequency intervals, and we roved the level of each stimulus over a large range (12 dB). Under these conditions, the DLF enhancement was still observed in all patients; this demonstrates that the effect cannot be explained simply by loudness cues. We then screened the patients for SOAEs to test whether the DLF enhancement effect could be explained by the presence of such emissions in the vicinity of the Fc. None of the patients exhibited SOAEs. Finally, we tested whether the patients had cochlear dead regions, i.e. regions lacking functional inner hair cells and/or auditory nerve fibres. Using a refined version of a non-invasive clinical test for the identification of dead regions, we assessed the presence of such regions in fine frequency steps (1/4 octave) up to very high frequencies. All of the patients had cochlear dead regions. The first two findings support the hypothesis that DLF enhancement is due to injury-induced central reorganization in the auditory system. The last one is consistent with neurophysiological data in animals, which suggest that complete deprivation from auditory input at certain cochlear sites may be a necessary condition for the occurrence of injury-induced cortical reorganization.

Aged↗

The neurophysiological basis of the auditory continuity illusion: a mismatch negativity study.

A sound turned off for a short moment can be perceived as continuous if the silent gap is filled with noise. The neural mechanisms underlying this "continuity illusion" were investigated using the mismatch negativity (MMN), an event-related potential reflecting the perception of a sudden change in an otherwise regular stimulus sequence. The MMN was recorded in four conditions using an oddball paradigm. The standards consisted of 500-Hz, 120-msec tone pips that were either physically continuous (Condition 1) or were interrupted by a 40-msec silent gap (Condition 2). The deviants consisted of the interrupted tone, but with the silent gap filled by a burst of bandpass-filtered noise. The noise either occupied the same frequency region as the tone and elicited the continuity illusion (Conditions 1a and 2a), or occupied a remote frequency region and did not elicit the illusion (Conditions 1b and 2b). We predicted that, if the continuity illusion is determined before MMN generation, then, other things being equal, the MMN should be larger in conditions where the deviants are perceived as continuous and the standards as interrupted or vice versa, than when both were perceived as continuous or both interrupted. Consistent with this prediction, we observed an interaction between standard type and noise frequency region, with the MMN being larger in Condition 1a than in Condition 1b, but smaller in Condition 2a than in Condition 2b. Because the subjects were instructed to ignore the tones and watch a silent movie during the recordings, the results indicate that the continuity illusion can occur outside the focus of attention. Furthermore, the latency of the MMN (less than approximately 200 msec postdeviance onset) places an upper limit on the stage of neural processing responsible for the illusion.

Acoustic Stimulation↗

Learning in discrimination of frequency or modulation rate: generalization to fundamental frequency discrimination.

Fifteen initially inexperienced subjects were trained for 4 weeks (12 2-h sessions) in frequency discrimination with pure tones around 88, 250, or 1605 Hz, or amplitude modulation rate discrimination of noise bands, using modulation rates around 88 or 250 Hz. Before, in the middle of, and after this training period, pure-tone frequency discrimination thresholds (DLFs), harmonic complex tone fundamental frequency discrimination thresholds (DLF0s), and amplitude modulation rate discrimination thresholds (DLFMs) were measured in several conditions including the trained one. Training in pure-tone frequency discrimination resulted in significantly larger improvements in DLF0s when the test complexes contained resolved harmonics than when they were composed of unresolved harmonics. This result supports the hypothesis that the discrimination of the F0 of resolved harmonics shares common underlying mechanisms with the frequency discrimination of pure tones. Training in rate discrimination did not result in larger DLF0 improvements for unresolved than for resolved harmonics.

Acoustic Stimulation↗

Neurodynamics for auditory stream segregation: tracking sounds in the mustached bat's natural environment.

During navigation and the search phase of foraging, mustached bats emit approximately 25 ms long echolocation pulses (at 10-40 Hz) that contain multiple harmonics of a constant frequency (CF) component followed by a short (3 ms) downward frequency modulation. In the context of auditory stream segregation, therefore, bats may either perceive a coherent pulse-echo sequence (PEPE...), or segregated pulse and echo streams (P-P-P... and E-E-E...). To identify the neural mechanisms for stream segregation in bats, we developed a simple yet realistic neural network model with seven layers and 420 nodes. Our model required recurrent and lateral inhibition to enable output nodes in the network to 'latch-on' to a single tone (corresponding to a CF component in either the pulse or echo), i.e., exhibit differential suppression by the alternating two tones presented at a high rate (> 10 Hz). To test the applicability of our model to echolocation, we obtained neurophysiological data from the primary auditory cortex of awake mustached bats. Event-related potentials reliably reproduced the latching behaviour observed at output nodes in the network. Pulse as well as nontarget (clutter) echo CFs facilitated this latching. Individual single unit responses were erratic, but when summed over several recording sites, they also exhibited reliable latching behaviour even at 40 Hz. On the basis of these findings, we propose that a neural correlate of auditory stream segregation is present within localized synaptic activity in the mustached bat's auditory cortex and this mechanism may enhance the perception of echolocation sounds in the natural environment.

Acoustic Stimulation↗

Local improvement in auditory frequency discrimination is associated with hearing-loss slope in subjects with cochlear damage.

Earlier data in the literature have shown local improvements in frequency discrimination performance near the cut-off frequency of steeply sloping, high-frequency hearing loss in subjects with cochlear damage. The general objective of the present study was to characterize further the relationships between this effect and various audiometric variables: namely, the slope, extent and shape of the hearing loss. In particular, we were interested in determining whether the effect was present in subjects with more moderately sloping hearing loss and/or other patterns of loss. Frequency difference limens (DLFs) were measured in 20 subjects (eight female, 12 male, median age 55.5 years) with high-frequency hearing loss. At least 12 frequencies were tested at intervals of 1/8 octave over a range of 1.5 octaves around the cut-off frequency for hearing loss (Fc). The Fc corresponded to the audiogram edge frequency and was defined as the highest test frequency, at the beginning of the slope, with a hearing threshold of no more than 5 dB HL above that of the best hearing frequency. The level of the test tones was randomized over a range of 6 dB around a nominal level, following an equal-loudness contour curve measured at 1/2-octave intervals. Results showed that DLFs were significantly smaller in a frequency band 1/4 octave wide centred on Fc than in the other bands. Furthermore, the average DLF measured in this band proved to be negatively correlated with the slope of hearing loss. No such significant relationship was found with the other audiometric indices considered, namely, the extent and maximum amount of hearing loss and the log-transformed cut-off frequency. The 20 subjects were divided into three groups according to the slope of their hearing loss relative to Fc (steep, >25 dB/1/2 octave; medium, between 12 and 25 dB/1/2 octave; and shallow, <12 dB/1/2 octave). A local improvement in DLF around Fc was observed in the steep- and medium-slope groups and was confirmed statistically in the steep-slope group. Similar measurements in subjects with low-frequency or notched hearing loss allowed us to establish the presence of similar local improvements in DLFs around audiogram edges. These results, which suggest the slope of the hearing loss to be the most important factor for the occurrence of local DLF improvements, are consistent with both an interpretation in terms of peripheral mechanisms and one in terms of central mechanisms, i.e. injury-induced neural reorganization.

Acoustic Stimulation↗

Auditory stream segregation on the basis of amplitude-modulation rate.

In this study, auditory stream segregation based on differences in the rate of envelope fluctuations--in the absence of spectral and temporal fine structure cues--was tested. The temporal sequences to segregate were composed of fully amplitude-modulated (AM) bursts of broadband noises A and B. All sequences were built by the reiteration of a ABA triplet where A modulation rate was fixed at 100 Hz and B modulation rate was variable. The first experiment was devoted to measuring the threshold difference in AM rate leading subjects to perceive the sequence as two streams as opposed to just one. The results of this first experiment revealed that subjects generally perceived the sequences as a single perceptual stream when the difference in AM rate between the A and B noises was smaller than 0.75 oct, and as two streams when the difference was larger than about 1.00 oct. These streaming thresholds were found to be substantially larger than, and not related to, the subjects' modulation-rate discrimination thresholds. The results of a second experiment demonstrated that AM-rate-based streaming was adversely affected by decreases in AM depth, but that segregation remained possible as long as the AM of either the A or B noises was above the subject's AM-detection threshold. The results of a third experiment indicated that AM-rate-based streaming effects were still observed when the modulations applied to the A and B noises were set individually, either at a constant level in dB above AM-detection threshold, or at levels at which they were of the same perceived strength. This finding suggests that AM-rate-based streaming is not necessarily mediated by perceived differences in AM depth. Altogether, the results of this study indicate that sequential sounds can be segregated on the sole basis of differences in the rate of their temporal fluctuations in the absence of other temporal or spectral cues.

Adult↗

Loudness changes associated with the perception of an auditory after-image.

The Zwicker tone (ZT) is an auditory sensation that occurs following the presentation of broadband noise containing a spectral notch. The present study aimed to test whether the changes in auditory thresholds that have been shown to follow the presentation of the ZT inducer are accompanied by suprathreshold effects. Using an interaural loudness-balance procedure, the loudness of probe tones presented after notched and after flat noise was compared. The results revealed small differences in the influence of the two types of noise on loudness at low intensities only. This suggests that the influence of notched noise stimulation on the auditory system is mediated by changes in the internal noise in auditory centres.

Adult↗

Evidence for two pitch encoding mechanisms using a selective auditory training paradigm.

The neural mechanisms underlying the perception of pitch, a sensory attribute of paramount importance in hearing, have been a matter of debate for over a century. A question currently at the heart of the debate is whether the pitch of all harmonic complex tones can be determined by the auditory system's using a single mechanism, or whether two different neural mechanisms are involved, depending on the stimulus conditions. When the harmonics are widely spaced, as is the case at high fundamental frequencies (FOs), and/or when the frequencies of the harmonics are low, the frequency components of the sound fall in different peripheral auditory channels and are then "resolved" by the peripheral auditory system. In contrast, at low F0s, or when the harmonics are high in frequency, several harmonics interact within the passbands of the same auditory filters, being thus "unresolved" by the peripheral auditory system. The idea that more than one mechanism mediates the encoding of pitch depending on the resolvability status of the harmonics was investigated here by testing for transfer of learning in F0 discrimination between different stimulus conditions involving either resolved or unresolved harmonics after specific training in one of these conditions. The results, which show some resolvability-specificity of F0-discrimination learning, support the hypothesis that two different underlying mechanisms mediate the encoding of the F0 of resolved and unresolved harmonics.

Adult↗