On musical interval perception for complex tones at very high frequencies.

Listeners appear able to extract a residue pitch from high-frequency harmonics for which phase locking to the temporal fine structure is weak or absent. The present study investigated musical interval perception for high-frequency harmonic complex tones using the same stimuli as Lau, Mehta, and Oxenham [J. Neurosci. 37, 9013-9021 (2017)]. Nine young musically trained listeners with especially good high-frequency hearing adjusted various musical intervals using harmonic complex tones containing harmonics 6-10. The reference notes had fundamental frequencies (F0s) of 280 or 1400 Hz. Interval matches were possible, albeit markedly worse, even when all harmonic frequencies were above the presumed limit of phase locking. Matches showed significantly larger systematic errors and higher variability, and subjects required more trials to finish a match for the high than for the low F0. Additional absolute pitch judgments from one subject with absolute pitch, for complex tones containing harmonics 1-5 or 6-10 with a wide range of F0s, were perfect when the lowest frequency component was below about 7 kHz, but at least 50% of responses were incorrect when it was 8 kHz or higher. The results are discussed in terms of the possible effects of phase-locking information and familiarity with high-frequency stimuli on pitch

Pitch perception at very high frequencies: On psychometric functions and integration of frequency information.

Lau et al. [J. Neurosci. 37, 9013-9021 (2017)] showed that discrimination of the fundamental frequency (F0) of complex tones with components in a high-frequency region was better than predicted from the optimal combination of information from the individual harmonics. The predictions depend on the assumption that psychometric functions for frequency discrimination have a slope of 1 at high frequencies. This was tested by measuring psychometric functions for F0 discrimination and frequency discrimination. Difference limens for F0 (F0DLs) and difference limens for frequency for each frequency component were also measured. Complex tones contained harmonics 6-10 and had F0s of 280 or 1400 Hz. Thresholds were measured using 210-ms tones presented diotically in diotic threshold-equalizing noise (TEN), and 1000-ms tones presented diotically in dichotic TEN. The slopes of the psychometric functions were close to 1 for all frequencies and F0s. The ratio of predicted to observed F0DLs was around 1 or smaller for both F0s, i.e., not super-optimal, and was significantly smaller for the low than for the high F0. The results are consistent with the idea that place information alone can convey pitch, but pitch is more salient when phase-locking information is available

Effect of Context on the Contribution of Individual Harmonics to Residue Pitch.

There is evidence that the contribution of a given harmonic in a complex tone to residue pitch is influenced by the accuracy with which the frequency of that harmonic is encoded. The present study investigated whether listeners adjust the weights assigned to individual harmonics based on acquired knowledge of the reliability of the frequency estimates of those harmonics. In a two-interval forced-choice task, seven listeners indicated which of two 12-harmonic complex tones had the higher overall pitch. In context trials (60 % of all trials), the fundamental frequency (F0) was 200 Hz in one interval and 200 + ΔF0 Hz in the other. In different (blocked) conditions, either the 3rd or the 4th harmonic (plus the 7th, 9th, and 12th harmonics), were replaced by narrowband noises that were identical in the two intervals. Feedback was provided. In randomly interspersed test trials (40 % of all trials), the fundamental frequency was 200 + ΔF0/2 Hz in both intervals; in the second interval, either the third or the fourth harmonic was shifted slightly up or down in frequency with equal probability. There were no narrowband noises. Feedback was not provided. The results showed that substitution of a harmonic by noise in context trials reduced the contribution of that harmonic to pitch judgements in the test trials by a small but significant amount. This is consistent with the notion that listeners give smaller weight to a harmonic or frequency region when they have learned that this frequency region does not provide reliable information for a given task

Specificity of the human frequency following response for carrier and modulation frequency assessed using adaptation

The frequency following response (FFR) is a scalp-recorded measure of phase-locked brainstem activity to stimulus-related periodicities. Three experiments investigated the specificity of the FFR for carrier and modulation frequency using adaptation. FFR waveforms evoked by alternating-polarity stimuli were averaged for each polarity and added, to enhance envelope, or subtracted, to enhance temporal fine structure information. The first experiment investigated peristimulus adaptation of the FFR for pure and complex tones as a function of stimulus frequency and fundamental frequency (F0). It showed more adaptation of the FFR in response to sounds with higher frequencies or F0s than to sounds with lower frequency or F0s. The second experiment investigated tuning to modulation rate in the FFR. The FFR to a complex tone with a modulation rate of 213 Hz was not reduced more by an adaptor that had the same modulation rate than by an adaptor with a different modulation rate (90 or 504 Hz), thus providing no evidence that the FFR originates mainly from neurons that respond selectively to the modulation rate of the stimulus. The third experiment investigated tuning to audio frequency in the FFR using pure tones. An adaptor that had the same frequency as the target (213 or 504 Hz) did not generally reduce the FFR to the target more than an adaptor that differed in frequency (by 1.24 octaves). Thus, there was no evidence that the FFR originated mainly from neurons tuned to the frequency of the target. Instead, the results are consistent with the suggestion that the FFR for low-frequency pure tones at medium to high levels mainly originates from neurons tuned to higher frequencies. Implications for the use and interpretation of the FFR are discussed

The binaural masking level difference: cortical correlates persist despite severe brain stem atrophy in progressive supranuclear palsy.

Under binaural listening conditions, the detection of target signals within background masking noise is substantially improved when the interaural phase of the target differs from that of the masker. Neural correlates of this binaural masking level difference (BMLD) have been observed in the inferior colliculus and temporal cortex, but it is not known whether degeneration of the inferior colliculus would result in a reduction of the BMLD in humans. We used magnetoencephalography to examine the BMLD in 13 healthy adults and 13 patients with progressive supranuclear palsy (PSP). PSP is associated with severe atrophy of the upper brain stem, including the inferior colliculus, confirmed by voxel-based morphometry of structural MRI. Stimuli comprised in-phase sinusoidal tones presented to both ears at three levels (high, medium, and low) masked by in-phase noise, which rendered the low-level tone inaudible. Critically, the BMLD was measured using a low-level tone presented in opposite phase across ears, making it audible against the noise. The cortical waveforms from bilateral auditory sources revealed significantly larger N1m peaks for the out-of-phase low-level tone compared with the in-phase low-level tone, for both groups, indicating preservation of early cortical correlates of the BMLD in PSP. In PSP a significant delay was observed in the onset of the N1m deflection and the amplitude of the P2m was reduced, but these differences were not restricted to the BMLD condition. The results demonstrate that although PSP causes subtle auditory deficits, binaural processing can survive the presence of significant damage to the upper brain stem.This work has been supported by the Wellcome Trust (Grants 088324 and 088263); Medical Research Council (G0700503 to B. C. P. Ghosh); Guarantors of Brain (to B. C. P. Ghosh); Raymond and Beverley Sackler Trust (to B. C. P. Ghosh); and National Institute of Health Research Cambridge Comprehensive Biomedical Research Centre including the CambridgeBrain Bank.This is the final version of the article. It first appeared from American Physiological Society via http://dx.doi.org/10.1152/jn.00062.201

Teologija na tržištu

One task intended to measure sensitivity to temporal fine structure (TFS) involves the discrimination of a harmonic complex tone from a tone in which all harmonics are shifted upwards by the same amount in hertz. Both tones are passed through a fixed bandpass filter centered on the high harmonics to reduce the availability of excitation-pattern cues and a background noise is used to mask combination tones. The role of frequency selectivity in this "TFS1" task was investigated by varying level. Experiment 1 showed that listeners performed more poorly at a high level than at a low level. Experiment 2 included intermediate levels and showed that performance deteriorated for levels above about 57 dB sound pressure level. Experiment 3 estimated the magnitude of excitation-pattern cues from the variation in forward masking of a pure tone as a function of frequency shift in the complex tones. There was negligible variation, except for the lowest level used. The results indicate that the changes in excitation level at threshold for the TFS1 task would be too small to be usable. The results are consistent with the TFS1 task being performed using TFS cues, and with frequency selectivity having an indirect effect on performance via its influence on TFS cues. (C) 2015 Acoustical Society of America

Combination of Spectral and Binaurally Created Harmonics in a Common Central Pitch Processor

A fundamental attribute of human hearing is the ability to extract a residue pitch from harmonic complex sounds such as those produced by musical instruments and the human voice. However, the neural mechanisms that underlie this processing are unclear, as are the locations of these mechanisms in the auditory pathway. The ability to extract a residue pitch corresponding to the fundamental frequency from individual harmonics, even when the fundamental component is absent, has been demonstrated separately for conventional pitches and for Huggins pitch (HP), a stimulus without monaural pitch information. HP is created by presenting the same wideband noise to both ears, except for a narrowband frequency region where the noise is decorrelated across the two ears. The present study investigated whether residue pitch can be derived by combining a component derived solely from binaural interaction (HP) with a spectral component for which no binaural processing is required. Fifteen listeners indicated which of two sequentially presented sounds was higher in pitch. Each sound consisted of two “harmonics,” which independently could be either a spectral or a HP component. Component frequencies were chosen such that the relative pitch judgement revealed whether a residue pitch was heard or not. The results showed that listeners were equally likely to perceive a residue pitch when one component was dichotic and the other was spectral as when the components were both spectral or both dichotic. This suggests that there exists a single mechanism for the derivation of residue pitch from binaurally created components and from spectral components, and that this mechanism operates at or after the level of the dorsal nucleus of the lateral lemniscus (brainstem) or the inferior colliculus (midbrain), which receive inputs from the medial superior olive where temporal information from the two ears is first combined

The Frequency Following Response (FFR) May Reflect Pitch-Bearing Information But is Not a Direct Representation of Pitch

The frequency following response (FFR), a scalp-recorded measure of phase-locked brainstem activity, is often assumed to reflect the pitch of sounds as perceived by humans. In two experiments, we investigated the characteristics of the FFR evoked by complex tones. FFR waveforms to alternating-polarity stimuli were averaged for each polarity and added, to enhance envelope, or subtracted, to enhance temporal fine structure information. In experiment 1, frequency-shifted complex tones, with all harmonics shifted by the same amount in Hertz, were presented diotically. Only the autocorrelation functions (ACFs) of the subtraction-FFR waveforms showed a peak at a delay shifted in the direction of the expected pitch shifts. This expected pitch shift was also present in the ACFs of the output of an auditory nerve model. In experiment 2, the components of a harmonic complex with harmonic numbers 2, 3, and 4 were presented either to the same ear (“mono”) or the third harmonic was presented contralaterally to the ear receiving the even harmonics (“dichotic”). In the latter case, a pitch corresponding to the missing fundamental was still perceived. Monaural control conditions presenting only the even harmonics (“2 + 4”) or only the third harmonic (“3”) were also tested. Both the subtraction and the addition waveforms showed that (1) the FFR magnitude spectra for “dichotic” were similar to the sum of the spectra for the two monaural control conditions and lacked peaks at the fundamental frequency and other distortion products visible for “mono” and (2) ACFs for “dichotic” were similar to those for “2 + 4” and dissimilar to those for “mono.” The results indicate that the neural responses reflected in the FFR preserve monaural temporal information that may be important for pitch, but provide no evidence for any additional processing over and above that already present in the auditory periphery, and do not directly represent the pitch of dichotic stimuli

Effect of diotic versus dichotic presentation on the pitch perception of tone complexes at medium and very high frequencies

Abstract Difference limens for fundamental frequency (F0), F0DLs, are usually small for complex tones containing low harmonics that are resolved in the auditory periphery, but worsen when the rank of the lowest harmonic increases above about 6–8 and harmonics become less resolved. The traditional explanation for this, in terms of resolvability, has been challenged and an alternative explanation in terms of harmonic rank was suggested. Here, to disentangle the effects of resolvability and harmonic rank the complex tones were presented either diotically (all harmonics to both ears) or dichotically (even and odd harmonics to opposite ears); the latter increases resolvability but does not affect harmonic rank. F0DLs were measured for 14 listeners for complex tones containing harmonics 6–10 with F0s of 280 and 1400 Hz, presented diotically or dichotically. For the low F0, F0DLs were significantly lower for the dichotic than for the diotic condition. This is consistent with a benefit of increased resolvability of harmonics for F0 discrimination and extends previous results to harmonics as low as the sixth. In contrast, for the high F0, F0DLs were similar for the two presentation modes, adding to evidence for differences in pitch perception between tones with low-to-medium and very-high frequency content