Hedwig E. Gockel

Properties of auditory stream formation

A sequence of sounds may be heard as coming from a single source (called fusion or coherence) or from two or more sources (called fission or stream segregation). Each perceived source is called a ‘stream’. When the differences between successive sounds are very large, fission nearly always occurs, whereas when the differences are very small, fusion nearly always occurs. When the differences are intermediate in size, the percept often ‘flips’ between one stream and multiple streams, a property called ‘bistability’. The flips do not generally occur regularly in time. The tendency to hear two streams builds up over time, but can be partially or completely reset by a sudden change in the properties of the sequence or by switches in attention. Stream formation depends partly on the extent to which successive sounds excite different ‘channels’ in the peripheral auditory system. However, other factors can play a strong role; multiple streams may be heard when successive sounds are presented to the same ear and have essentially identical excitation patterns in the cochlea. Differences between successive sounds in temporal envelope, fundamental frequency, phase spectrum and lateralization can all induce a percept of multiple streams. Regularities in the temporal pattern of elements within a stream can help in stabilizing that stream.

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.

Subcortical neural synchrony and absolute thresholds predict frequency discrimination independently

The neural mechanisms of pitch coding have been debated for more than a century. The two main mechanisms are coding based on the profiles of neural firing rates across auditory nerve fibers with different characteristic frequencies (place-rate coding), and coding based on the phase-locked temporal pattern of neural firing (temporal coding). Phase locking precision can be partly assessed by recording the frequency-following response (FFR), a scalp-recorded electrophysiological response that reflects synchronous activity in subcortical neurons. Although features of the FFR have been widely used as indices of pitch coding acuity, only a handful of studies have directly investigated the relation between the FFR and behavioral pitch judgments. Furthermore, the contribution of degraded neural synchrony (as indexed by the FFR) to the pitch perception impairments of older listeners and those with hearing loss is not well known. Here, the relation between the FFR and pure-tone frequency discrimination was investigated in listeners with a wide range of ages and absolute thresholds, to assess the respective contributions of subcortical neural synchrony and other age-related and hearing loss-related mechanisms to frequency discrimination performance. FFR measures of neural synchrony and absolute thresholds independently contributed to frequency discrimination performance. Age alone, i.e., once the effect of subcortical neural synchrony measures or absolute thresholds had been partialed out, did not contribute to frequency discrimination. Overall, the results suggest that frequency discrimination of pure tones may depend both on phase locking precision and on separate mechanisms affected in hearing loss.

Asymmetry of masking between complex tones and noise: The role of temporal structure and peripheral compression

Thresholds for the detection of harmonic complex tones in noise were measured as a function of masker level. The rms level of the masker ranged from 40 to 70 dB SPL in 10-dB steps. The tones had a fundamental frequency (F0) of 62.5 or 250 Hz, and components were added in either cosine or random phase. The complex tones and the noise were bandpass filtered into the same frequency region, from the tenth harmonic up to 5 kHz. In a different condition, the roles of masker and signal were reversed, keeping all other parameters the same; subjects had to detect the noise in the presence of a harmonic tone masker. In both conditions, the masker was either gated synchronously with the 700-ms signal, or it started 400 ms before and stopped 200 ms after the signal. The results showed a large asymmetry in the effectiveness of masking between the tones and noise. Even though signal and masker had the same bandwidth, the noise was a more effective masker than the complex tone. The degree of asymmetry depended on F0, component phase, and the level of the masker. The maximum difference between masked thresholds for tone and noise was about 28 dB; this occurred when the F0 was 62.5 Hz, the components were in cosine phase, and the masker level was 70 dB SPL. In most conditions, the growth-of-masking functions had slopes close to 1 (on a dB versus dB scale). However, for the cosine-phase tone masker with an F0 of 62.5 Hz, a 10-dB increase in masker level led to an increase in masked threshold of the noise of only 3.7 dB, on average. We suggest that the results for this condition are strongly affected by the active mechanism in the cochlea.

Louder sounds can produce less forward masking: Effects of component phase in complex tones

The influence of the degree of envelope modulation and periodicity on the loudness and effectiveness of sounds as forward maskers was investigated. In the first experiment, listeners matched the loudness of complex tones and noise. The tones had a fundamental frequency (F0) of 62.5 or 250 Hz and were filtered into a frequency range from the 10th harmonic to 5000 Hz. The Gaussian noise was filtered in the same way. The components of the complex tones were added either in cosine phase (CPH), giving a large crest factor, or in random phase (RPH), giving a smaller crest factor. For each F0, subjects matched the loudness between all possible stimulus pairs. Six different levels of the fixed stimulus were used, ranging from about 30 dB SPL to about 80 dB SPL in 10-dB steps. Results showed that, at a given overall level, the CPH and the RPH tones were louder than the noise, and that the CPH tone was louder than the RPH tone. The difference in loudness was larger at medium than at low levels and was only slightly reduced by the addition of a noise intended to mask combination tones. The differences in loudness were slightly smaller for the higher than for the lower F0. In the second experiment, the stimuli with the lower F0s were used as forward maskers of a 20-ms sinusoid, presented at various frequencies within the spectral range of the maskers. Results showed that the CPH tone was the least effective forward masker, even though it was the loudest. The differences in effectiveness as forward maskers depended on masker level and signal frequency; in order to produce equal masking, the level of the CPH tone had to be up to 35 dB above that of the RPH tone and the noise. The implications of these results for models of loudness are discussed and a model is presented based on neural activity patterns in the auditory nerve; this predicts the general pattern of loudness matches. It is suggested that the effects observed in the experiments may have been influenced by two factors: cochlear compression and suppression.

Influence of rate of change of frequency on the overall pitch of frequency-modulated tones

The mechanism(s) determining pitch may assign less weight to portions of a sound where the frequency is changing rapidly. The present experiments explored the possible effect of this on the overall pitch of frequency-modulated sounds. Pitch matches were obtained between an adjustable unmodulated sinusoid and a sinusoidal carrier that was frequency modulated using a highly asymmetric function with the form of a repeating U or inverted U shaped function. The amplitude was constant during the 400-ms presentation time of each stimulus, except for 10-ms raised-cosine onset and offset ramps. In experiment 1, the carrier level was 50 dB SPL and the geometric mean of the instantaneous frequency of the modulated carrier, fc, was either 0.5, 1, 2, or 8 kHz. The modulation rate (fm) was 5, 10, or 20 Hz. The overall depth (maximum to minimum) of the FM was 8% of fc. For all carrier frequencies, the matched frequency was shifted away from the mean carrier frequency, downwards for the U shaped function stimuli and upwards for the repeated inverted U shaped function stimuli. The shift was typically slightly greater than 1% of fc, and did not vary markedly with fc. The effect of fm was small, but there was a trend for the shifts to decrease with increasing fm for fc = 0.5 kHz and to increase with increasing fm for fc = 2 kHz. In experiment 2, the carrier level was reduced to 20 dB SL and matches were obtained only for fc = 2 kHz. Shifts in matched frequency of about 1% were still observed, but the trend for the shifts to increase with increasing fm no longer occurred. In experiment 3, matches were obtained for a 4-kHz carrier at 50 dB SPL. Shifts of about 1% again occurred, which did not vary markedly with fm. The shifts in matched frequency observed in all three experiments are not predicted by models based on the amplitude- or intensity-weighted average of instantaneous frequency (EWAIF or IWAIF). The shifts (and the pitch shifts observed earlier for two-tone complexes and for stimuli with simultaneous AM and FM) are consistent with a model based on the assumption that the overall pitch of a frequency-modulated sound is determined from a weighted average of period estimates, with the weight attached to a given estimate being inversely related to the short-term rate of change of period and directly related to a compressive function of the amplitude.

Effect of duration on the frequency discrimination of individual partials in a complex tone and on the discrimination of fundamental frequency

Thresholds for the discrimination of fundamental frequency (FODLs) and frequency difference limens (FDLs) for individual partials within a complex tone (F0=250 Hz, harmonics 1-7) were measured for stimulus durations of 200, 50, and 16 ms. The FDLs increased with decreasing duration. Although the results differed across subjects, the effect of duration generally decreased as the harmonic number increased from 1 to 4, then increased as the harmonic number increased to 6, and finally decreased for the seventh harmonic. For each duration, FODLs were smaller than the smallest FDL for any individual harmonic, indicating that information is combined across harmonics in the discrimination of FO. FODLs predicted from the FDLs corresponded well with observed FODLs for the 200- and 16-ms durations but were significantly larger than observed FODLs for the 50-ms duration. A supplementary pitch-matching experiment using two subjects indicated that the contribution of the seventh harmonic to the pitch of the 16-ms complex tone was smaller than would be predicted from the FDL for that harmonic. The results are consistent with the idea that the dominant region shifts upward with decreasing duration, but that the weight assigned to individual harmonics is not always adjusted in an optimal way.

Dominance region for pitch: effects of duration and dichotic presentation.

The dominance region (DR) for pitch was determined for 16- and 200-ms complex tones containing the first seven harmonics of a fundamental frequency (F0) of 250 Hz. A tone was presented with one of the harmonics mistuned upwards or downwards by 3%, followed 500 ms later by a perfectly harmonic tone of the same duration. Listeners adjusted the F0 of the harmonic tone so that its pitch matched that of the mistuned complex. In experiment 1, stimuli were presented monaurally. The DR was significantly higher in harmonic number for the short than for the long duration. The overall sum of the pitch shifts produced by all harmonics was significantly larger for the short than for the long duration, presumably due to stronger perceptual fusion for the former. In experiment 2, the mistuned harmonic was presented only contralaterally to the remainder of the complex. A similar shift in the DR with duration was observed, although the pitch shifts were smaller than for monaural presentation. There was no significant effect of duration on the overall pitch shifts. The results are discussed in terms of pattern recognition and autocorrelation models of pitch perception, and a role of attention in pitch matching is suggested.

Frequency discrimination duration effects for Huggins pitch and narrowband noise (L)

Frequency difference limens (FDLs) were measured for Huggins pitch (HP) stimuli, consisting of a 30-Hz wide band of interaurally decorrelated noise in a diotic low-pass noise and for 30-Hz wide bands of diotic narrowband noise presented in a diotic low-pass noise background. FDLs at a 400-ms duration for the two stimulus types were equated by adjusting the level of the narrowband noise relative to the background. The effects of duration on the FDLs were then measured for center frequencies of 300, 600, and 900 Hz. Although the results were compromised by floor effects at 900 Hz, at 300 and 600 Hz, the duration effects were very similar for the HP and narrowband noise stimuli, with a large improvement in performance between 100 and 400 ms. In contrast to previous results for pure tones, the effect of duration was independent of frequency. The results suggest that: (1) Binaural and monaural pitches may be processed using a common mechanism; (2) discrimination performance for HP and low-sensation-level narrowband noise stimuli is not determined by the number of waveform periods.

Asymmetry of masking between complex tones and noise: Partial loudness

This experiment examined the partial masking of periodic complex tones by a background of noise, and vice versa. The tones had a fundamental frequency (F0) of 62.5 or 250 Hz, and components were added in either cosine phase (CPH) or random phase (RPH). The tones and the noise were bandpass filtered into the same frequency region, from the tenth harmonic up to 5 kHz. The target alone was alternated with the target and the background; for the mixture, the background and target were either gated together, or the background was turned on 400 ms before, and off 200 ms after, the target. Subjects had to adjust the level of either the target alone or the target in the background so as to match the loudness of the target in the two intervals. The overall level of the background was 50 dB SPL, and loudness matches were obtained for several fixed levels of the target alone or in the background. The resulting loudness-matching functions showed clear asymmetry of partial masking. For a given target-to-background ratio, the partial loudness of a complex tone in a noise background was lower than the partial loudness of a noise in a complex tone background. Expressed as the target-to-background ratio required to achieve a given loudness, the asymmetry typically amounted to 12-16 dB. When the F0 of the complex tone was 62.5 Hz, the asymmetry of partial masking was greater for CPH than for RPH. When the F0 was 250 Hz, the asymmetry was greater for RPH than for CPH. Masked thresholds showed the same pattern as for partial masking for both F0s. Onset asynchrony had some effect on the loudness matching data when the target was just above its masked threshold, but did not significantly affect the level at which the target in the background reached its unmasked loudness. The results are interpreted in terms of the temporal structure of the stimuli.