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Functional role of auditory cortex in frequency processing and pitch perception.

Microelectrode studies in nonhuman primates and other mammals have demonstrated that many neurons in auditory cortex are excited by pure tone stimulation only when the tone's frequency lies within a narrow range of the audible spectrum. However, the effects of auditory cortex lesions in animals and humans have been interpreted as evidence against the notion that neuronal frequency selectivity is functionally relevant to frequency discrimination. Here we report psychophysical and anatomical evidence in favor of the hypothesis that fine-grained frequency resolution at the perceptual level relies on neuronal frequency selectivity in auditory cortex. An adaptive procedure was used to measure difference thresholds for pure tone frequency discrimination in five humans with focal brain lesions and eight normal controls. Only the patient with bilateral lesions of primary auditory cortex and surrounding areas showed markedly elevated frequency difference thresholds: Weber fractions for frequency direction discrimination ("higher"-"lower" pitch judgments) were about eightfold higher than Weber fractions measured in patients with unilateral lesions of auditory cortex, auditory midbrain, or dorsolateral frontal cortex; Weber fractions for frequency change discrimination ("same"-"different" pitch judgments) were about seven times higher. In contrast, pure-tone detection thresholds, difference thresholds for pure tone duration discrimination centered at 500 ms, difference thresholds for vibrotactile intensity discrimination, and judgments of visual line orientation were within normal limits or only mildly impaired following bilateral auditory cortex lesions. In light of current knowledge about the physiology and anatomy of primate auditory cortex and a review of previous lesion studies, we interpret the present results as evidence that fine-grained frequency processing at the perceptual level relies on the integrity of finely tuned neurons in auditory cortex.

Acoustic Stimulation↗

The slow formation of a pitch percept beyond the ending time of a short tone burst.

The discriminability of short tone bursts differing in frequency was measured in terms of the sensitivity index d' as a function of interstimulus interval (ISI). The two stimuli presented on each trial consisted of either 6 or 30 sinusoidal cycles. When the frequency of the first stimulus varied randomly and widely from trial to trial (Experiment 1), discriminability was maximal for an ISI of about 400 msec in the 6-cycles condition and for a significantly longer ISI (of about 1 sec) in the 30-cycles condition. However, when the first stimulus had only two possible frequencies and the second stimulus was fixed (Experiment 2), the optimal ISI appeared to be about 400 msec in both conditions. A final experiment confirmed that, for tone bursts of 30 cycles, the optimal ISI was dependent on the perceptual uncertainty of the first stimulus. These results support the idea that the duration required to perceive the pitch of a sound as accurately as possible may far exceed the duration of the stimulus itself. More importantly, they indicate that the required duration is not a constant.

Auditory Perception↗

Effects of onset asynchrony on pitch perception: adaptation or grouping?

A previous paper by Darwin and Ciocca [J. Acoust. Soc. Am. 91, 3381-3390 (1992)] showed that a slightly mistuned frequency component of a (target) harmonic complex produced smaller pitch shifts in the target if it started 160 ms or more before the other components than if all the components were simultaneous. Three experiments investigated whether this effect of onset asynchrony is due to peripheral adaptation to the leading portion of the mistuned component or to perceptual grouping. The first two experiments showed that the effect of asynchrony could be influenced by grouping mechanisms without changing the amount of adaptation produced by the leading portion of the mistuned component. In the first experiment, the effect of asynchrony was reduced by the presence of an additional (captor) complex which was harmonically related to the mistuned component and synchronous with just its leading portion. In experiment 2, the effect of asynchrony was increased by presenting a captor that was synchronous with the entire mistuned component. This capturing effect was independent of the harmonic relation between the captor and the mistuned component at 40-ms asynchrony; at 160 ms the effect of asynchrony increased further only if the captor and the mistuned component were harmonically related. In the third experiment, the expected amount of adaptation was increased (relative to that produced by a single sine precursor) by presenting several components that were close in frequency to the mistuned component and synchronous with its leading portion.(ABSTRACT TRUNCATED AT 250 WORDS)

Acoustic Stimulation↗

Perceived continuity and pitch perception.

Three experiments investigated the importance of perceived stimulus continuity for the perception of the fundamental frequency (F0) of an unresolved complex tone. The F0 of the complex was 250 Hz and the harmonics were bandpass filtered between 5500 and 7500 Hz. In the first experiment, F0 discrimination was measured for single-burst tones with durations of 20, 40, and 80 ms, and for stimuli containing two 20- or 40-ms tone bursts separated by an 8- or 16-ms gap. For the single-burst conditions, there was a large decrease in threshold as the duration was increased from 20 to 40 ms. However, performance in the gapped conditions was much worse than that for the single-burst condition with the same cumulative duration (e.g., two 20-ms bursts separated by 8 ms produced higher thresholds than one 40-ms burst). Adding a bandpass noise (with the same spectral envelope as the tone) in the gap between the two tone bursts improved performance to the level of the single-burst condition. When the noise was added, the two discrete tone bursts were perceived as one single tone burst interrupted by the noise, and this seemed to facilitate discrimination. In a second experiment, the effects on pitch of an envelope delay (phase shift) of 0.75 periods between two tone bursts separated by an 8-ms gap were investigated. If the gap was silent, the pitch of the pair was unaffected by the phase shift. However, if the gap contained the bandpass noise, the phase shift between the bursts did produce a significant downward shift in the pitch of the pair. Finally, the third experiment showed that presenting a noise before a single 20-ms burst may improve discrimination performance in some listeners, but not sufficiently to account for the results of the first experiment purely in terms of an improvement in the discriminability of the second tone burst in the pair. The experiments suggest that a level decrease between two tone bursts may disrupt or reset a long integration mechanism, decreasing performance. When there is no level decrease between the bursts, the auditory system may assume that the two bursts belong to the same single tone and analyze them together in order to derive F0.

Adult↗

The influence of duration on the perception of pitch in single and simultaneous complex tones.

The influence of duration on the virtual pitch of complex tones was measured using an absolute identification paradigm. If performance with two-tone complexes is expressed in terms of a single central frequency-coding noise function, this function is found to depend on duration in about the same way as the pure-tone difference limen function. The function is further found to be a reasonably good predictor of pitch identification performance with multitone complexes. Another experimental finding was that subjects tend to switch to the analytic mode of pitch perception when complex tones are shortened (i.e., they tend to hear the spectral pitches instead of the virtual ones). A third finding was that with simultaneous complex tones the degradation of each pitch percept depends not only on duration and harmonic order of the tone but also on the harmonic order of the other tone.

Audiometry↗

Dual temporal pitch percepts from acoustic and electric amplitude-modulated pulse trains.

Two experiments examined the perception of unmodulated and amplitude-modulated pulse trains by normally hearing listeners and cochlear implantees. Four normally hearing subjects listened to acoustic pulse trains, which were band-pass filtered between 3.9 and 5.3 kHz. Four cochlear implantees, all postlinguistically deaf users of the Mini System 22 implant, listened to current pulse trains produced at a single electrode position. In the first experiment, a set of nine loudness-balanced unmodulated stimuli with rates between 60 and 300 Hz were presented in a multidimensional scaling task. The resultant stimulus spaces for both subject groups showed a single dimension associated with the rate of the stimuli. In the second experiment, a set of ten loudness-balanced modulated stimuli was constructed, with carrier rates between 140 and 300 Hz, and modulation rates between 60 and 150 Hz. The modulation rates were integer submultiples of the carrier rates, and each modulation period consisted of one higher-intensity pulse and one or more identical lower-intensity pulses. The modulation depth of each stimulus was adjusted so that its pitch was judged to be higher or lower 50% of the time than that of an unmodulated pulse train having a rate equal to the geometric mean of the carrier and modulation rates. A multidimensional scaling task with these ten stimuli resulted in two-dimensional stimulus spaces, with dimensions corresponding to carrier and modulation rates. A further investigation with one normally hearing subject showed that the perceptual weighting of the two dimensions varied systematically with modulation depth. It was concluded that, when filtered appropriately, acoustic pulse trains can be used to produce percepts in normal listeners that share common features with those experienced by subjects listening through one channel of a cochlear implant, and that the central auditory system can extract two temporal patterns arising from the same cochlear location.

Acoustics↗

Periodicity pitch perception of retarded children.

In order to expand the body of information concerning auditory function in the presence of the generalized cortical dysfunction associated with severe mental retardation, the discrimination of periodicity pitch signals by nonverbal severely retarded children and adults of normal intelligence was examined using an operant conditioning adaptation of the psychophysical method of constant stimulus differences. The periodicity pitch difference limens (DLs) based on changes in repetition rate were compared with frequency DLs obtained from low (150 Hz) and high frequency (2200 Hz) sinusoids. Periodicity pitch signals were generated by gating a 2200-Hz sinusoid over a range from 141 to 159 pulses per sec and passing the signals through a narrow band-pass filter to ensure no low frequency energy was present. The DLs (75% correct discriminations) were computed from the three types of discrimination performance curves. Standard errors of the means were small and retarded subjects showed good test-retest reliability. The results indicated that the simple and complex pitch discrimination abilities for both normal and retarded subjects were similar.

Adult↗

Grouping in pitch perception: effects of onset asynchrony and ear of presentation of a mistuned component.

Three experiments investigated how the onset asynchrony and ear of presentation of a single mistuned frequency component influence its contribution to the pitch of an otherwise harmonic complex tone. Subjects matched the pitch of the target complex by adjusting the pitch of a second similar but strictly periodic complex tone. When the mistuned component (the 4th harmonic of a 155 Hz fundamental) started 160 ms or more before the remaining harmonics but stopped simultaneously with them, it made a reduced contribution to the pitch of the complex. It made no contribution if it started more than 300 ms before. Pitch shifts and their reduction with onset time were larger for short (90 ms) sounds than for long (410 ms). Pitch shifts were slightly larger when the mistuned component was presented to the same ear as the remaining 11 in-tune harmonics than to the opposite ear. Adding a "captor" complex tone with a fundamental of 200 Hz and a missing 3rd harmonic to the contralateral ear did not augment the effect of onset time, even though the captor was synchronous with the mistuned harmonic, the mistuned component was equal in frequency to the missing 3rd harmonic of the captor complex tone and it was played to the same ear as the captor. The results show that a difference in onset time can prevent a resolved frequency component from contributing to the pitch of a complex tone even though it is present throughout that complex tone.

Acoustic Stimulation↗