Aging and auditory temporal sequencing: ordering the elements of repeating tone patterns

In a series of experiments, we examined age-related differences in adults' ability to order sequences of tones presented at various speeds and in contexts designed to promote or to impede stream segregation. In Experiment 1, 32 listeners (16 young, 16 old) were required to identify two repeating sequences that consisted of four tones (two from a high and two from a low frequency range) in different order. In Experiment 2, 32 listeners were required to judge whether the two recycled patterns from Experiment 1 were the same or different. In Experiment 3, four young and four old listeners were tested on the tasks of Experiment 2 over an extended period. In Experiment 4, 16 young and 16 old listeners were tested with sequences that were not recycled and were composed of tones drawn from a narrow frequency range. Elderly adults were less able than young adults to distinguish between tone sequences with contrasting order, regardless of the speed of presentation, the nature of the task (identification vs. same/different), the amount of practice, the frequency separation of the tones, or the presence or absence of recycling. These findings provide evidence of a temporal sequencing impairment in elderly listeners but reveal no indication of age differences in streaming processes.     

Effects of the build-up and resetting of auditory stream segregation on temporal discrimination

The tendency to hear a tone sequence as 2 or more streams (segregated) builds up, but a sudden change in properties can reset the percept to 1 stream (integrated). This effect has not hitherto been explored using an objective measure of streaming. Stimuli comprised a 2.0-s fixed-frequency inducer followed by a 0.6-s test sequence of alternating pure tones (3 low [L]-high [H] cycles). Listeners compared intervals for which the test sequence was either isochronous or the H tones were slightly delayed. Resetting of segregation should make identifying the anisochronous interval easier. The HL frequency separation was varied (0-12 semitones), and properties of the inducer and test sequence were set to the same or different values. Inducer properties manipulated were frequency, number of onsets (several short bursts vs. one continuous tone), tone:silence ratio (short vs. extended bursts), level, and lateralization. All differences between the inducer and the L tones reduced temporal discrimination thresholds toward those for the no-inducer case, including properties shown previously not to affect segregation greatly. Overall, it is concluded that abrupt changes in a sequence cause resetting and improve subsequent temporal discrimination.     

Effects of context on auditory stream segregation

The authors examined the effect of preceding context on auditory stream segregation. Low tones (A), high tones (B), and silences (-) were presented in an ABA- pattern. Participants indicated whether they perceived 1 or 2 streams of tones. The A tone frequency was fixed, and the B tone was the same as the A tone or had 1 of 3 higher frequencies. Perception of 2 streams in the current trial increased with greater frequency separation between the A and B tones (Delta f). Larger Delta f in previous trials modified this pattern, causing less streaming in the current trial. This occurred even when listeners were asked to bias their perception toward hearing 1 stream or 2 streams. The effect of previous Delta f was not due to response bias because simply perceiving 2 streams in the previous trial did not cause less streaming in the current trial. Finally, the effect of previous ?f was diminished, though still present, when the silent duration between trials was increased to 5.76 s. The time course of this context effect on streaming implicates the involvement of auditory sensory memory or neural adaptation.     

Effect of rhythmic grouping on stream segregation

Three experiments investigated the role of temporal grouping on auditory stream segregation. For sounds that formed frequency streams (e.g., 400 Hz, 500 Hz, 1600 Hz, and 2000 Hz), the effect of rhythm was minimal. Temporal grouping did not affect judgments of stream segregation and did not affect difficulty of sequence identification. In contrast, for sounds that tended to form one coherent sequence (e.g., 750 Hz, 1500 Hz, 3000 Hz, and 6000 Hz), temporal grouping affected judgments of stream segregation as well as difficulty of identification. The temporal grouping could space the three lower or three higher pitch tones equally in time and this induced isochronous stream segregation. Subjects could not interleave the resulting streams, and identification became far more difficult. The role of rhythmic grouping is therefore contextual, depending on the relationships between the elements as well as the order of the elements.     

Effects of time intervals and tone durations on auditory stream segregation

Adult listeners rated the difficulty of hearing a single coherent stream in a sequence of high (H) and low (L) tones that alternated in a repetitive galloping pattern (HLH-HLH-HLH...). They could hear the gallop when the sequence was perceived as a single stream, but when it segregated into two substreams, they heard H-H- ... in one stream and L-L- ... in the other. The onset-to-onset time of the tones, their duration, the interstimulus interval (ISI) between tones of the same frequency, and the frequency separation between H and L tones were varied. Subjects' ratings on a 7-point scale showed that the well-known effect of speed's increasing stream segregation is primarily due to its effect on the ISI between tones in the same frequency region. This has implications for several theories of streaming.     

The effect of marker frequency disparity on the discrimination of gap duration in monkeys

Duration-discrimination thresholds of the silent interval (gap) between two successive tones (markers) were measured in four Japanese monkeys. The task was serial discrimination, and monkeys were required to release the lever when the gap duration decreased from 200 ms. Monkeys successfully acquired the task, and gap thresholds of monkeys were revealed to be larger than previous data with human subjects. Gap thresholds were not affected by marker frequency when the two markers were identical in frequency, though the thresholds increased when large frequency differences existed between markers. The effect of marker frequency disparity on gap thresholds in monkeys is discussed in terms of the difficulty in integrating information from discrete frequency channels.     

The effect of tone duration on auditory stream formation

In a study in which the effect of tone duration on the formation of auditory streams was investigated, subjects were presented with 15-sec alternating pure-tone sequences (ABAB …) and were asked to orient their attention over the duration of the sequence toward hearing either a temporally coherent or a segregated percept. At stimulus offset, the subjects indicated whether their percept at the end of the stimulus had been that of a temporally coherent ABAB trill or that of segregated A and B streams. The experimental results indicated that the occurrence of stream segregation increases as (1) the duration of the A and B tones increases in unison and (2) the difference in duration between the A and B tones increases, with the duration differences between the tones producing the strongest segregation effects. A comparison of these experimental results with those of other studies strongly suggests that the time interval between the offset and onset of consecutive tones in the same frequency range is the most important temporal factor affecting auditory stream formation. Furthermore, a simulation of the experimental results by the Beauvois and Meddis (1996) stream segregation model suggests that both the tone duration effects reported here and Gestalt auditory grouping on the basis of temporal proximity can be understood in terms of low-level neurophysiological processes and peripheral-channeling factors.     

Effects of similarity in bandwidth on the auditory sequential streaming of two-tone complexes

We investigated the perceptual grouping of sequentially presented sounds--auditory stream segregation. It is well established that sounds heard as more similar in quality, or timbre, are more likely to be grouped into the same auditory stream. However, it is often unclear exactly what acoustic factors determine timbre. In this study, we presented various sequences of simple sounds, each comprising two frequency components (two-tone complexes), and measured their perceptual grouping. We varied only one parameter between trials, the intercomponent separation for some of the complexes, and examined the effects on stream segregation. Four hypotheses are presented that might predict the extent of streaming. Specifically, least streaming might be expected when the sounds were most similar in either (1) the frequency regions in which they have energy (maximum spectral overlap), (2) their auditory bandwidths, (3) their relative bandwidths, or (4) the rate at which the two components beat together (intermodulation rate). It was found that least streaming occurred when sounds were most similar in either their auditory or their relative bandwidths. Although these two hypotheses could not be distinguished, the results were clearly different from those predicted by hypotheses (1) and (4). The implications for models of stream segregation are discussed.     

The relation between auditory temporal interval processing and sequential stream segregation examined with stimulus laterality differences

In this study, we examine the effects of laterality differences between noise bursts on two objective measures of temporal interval processing (gap detection and temporal asymmetry detection) and one subjective measure of temporal organization (stream segregation). Noise bursts were lateralized by presentation to different ears or dichotic presentation with oppositely signed interaural level (ILD) or time (ITD) differences. Objective thresholds were strongly affected by ear-of-entry differences, were moderately affected by ILD differences, but were unaffected by ITD differences. Subjectively, A and B streams segregated well on the basis of ear-of-entry or ILD differences but segregated poorly on the basis of ITD differences. These results suggest that perceptual segregation may be driven more effectively by differential activation of the two ears (peripheral channeling) than by differences in perceived laterality.     

Perceptual organization of complex auditory sequences: effect of number of simultaneous subsequences and frequency separation

Previous findings on streaming are generalized to sequences composed of more than 2 subsequences. A new paradigm identified whether listeners perceive complex sequences as a single unit (integrative listening) or segregate them into 2 (or more) perceptual units (stream segregation). Listeners heard 2 complex sequences, each composed of 1, 2, 3, or 4 subsequences. Their task was to detect a temporal irregularity within 1 subsequence. In Experiment 1, the smallest frequency separation under which listeners were able to focus on 1 subsequence was unaffected by the number of co-occurring subsequences; nonfocused sounds were not perceptually organized into streams. In Experiment 2, detection improved progressively, not abruptly, as the frequency separation between subsequences increased from 0.25 to 6 auditory filters. The authors propose a model of perceptual organization of complex auditory sequences.     

Segregated in perception,integrated for action: Immunity of rhythmic sensorimotor coordination to auditory stream segregation

Auditory stream segregation can occur when tones of different pitch (A, B) are repeated cyclically: The larger the pitch separation and the faster the tempo, the more likely perception of two separate streams is to occur. The present study assessed stream segregation in perceptual and sensorimotor tasks, using identical ABBABB … sequences. The perceptual task required detection of single phase-shifted A tones; this was expected to be facilitated by the presence of B tones unless segregation occurred. The sensorimotor task required tapping in synchrony with the A tones; here the phase correction response (PCR) to shifted A tones was expected to be inhibited by B tones unless segregation occurred. Two sequence tempi and three pitch separations (2, 10, and 48 semitones) were used with musically trained participants. Facilitation of perception occurred only at the smallest pitch separation, whereas the PCR was reduced equally at all separations. These results indicate that auditory action control is immune to perceptual stream segregation, at least in musicians. This may help musicians coordinate with diverse instruments in ensemble playing.     

Toward a neurophysiological theory of auditory stream segregation

Auditory stream segregation (or streaming) is a phenomenon in which 2 or more repeating sounds differing in at least 1 acoustic attribute are perceived as 2 or more separate sound sources (i.e., streams). This article selectively reviews psychophysical and computational studies of streaming and comprehensively reviews more recent neurophysiological studies that have provided important insights into the mechanisms of streaming. On the basis of these studies, segregation of sounds is likely to occur beginning in the auditory periphery and continuing at least to primary auditory cortex for simple cues such as pure-tone frequency but at stages as high as secondary auditory cortex for more complex cues such as periodicity pitch. Attention-dependent and perception-dependent processes are likely to take place in primary or secondary auditory cortex and may also involve higher level areas outside of auditory cortex. Topographic maps of acoustic attributes, stimulus-specific suppression, and competition between representations are among the neurophysiological mechanisms that likely contribute to streaming. A framework for future research is proposed.     

Sequential grouping of pure-tone percepts evoked by the segregation of components from a complex tone

A sudden change applied to a single component can cause its segregation from an ongoing complex tone as a pure-tone-like percept. Three experiments examined whether such pure-tone-like percepts are organized into streams by extending the research of Bregman and Rudnicky (1975). Those authors found that listeners struggled to identify the presentation order of 2 pure-tone targets of different frequency when they were flanked by 2 lower frequency "distractors." Adding a series of matched-frequency "captor" tones, however, improved performance by pulling the distractors into a separate stream from the targets. In the current study, sequences of discrete pure tones were substituted by sequences of brief changes applied to an otherwise constant 1.2-s complex tone. Pure-tone-like percepts were evoked by applying 6-dB increments to individual components of a complex comprising harmonics 1-7 of 300 Hz (Experiment 1) or 0.5-ms changes in interaural time difference to individual components of a log-spaced complex (range 160-905 Hz; Experiment 2). Results were consistent with the earlier study, providing clear evidence that pure-tone-like percepts are organized into streams. Experiment 3 adapted Experiment 1 by presenting a global amplitude increment either synchronous with, or just after, the last captor prior to the 1st distractor. In the former case, for which there was no pure-tone-like percept corresponding to that captor, the captor sequence did not aid performance to the same extent as previously. It is concluded that this change to the captor-tone stream partially resets the stream-formation process, and so the distractors and targets became likely to integrate once more.     

The perceived tempi of coherent and streaming tone sequences: II

Previous research demonstrated, by means of a reaction time technique, that there is no difference between the perceptual onset asynchronies (POAs) of nonalternating tone sequences and of frequency-alternating tone sequences, even if the latter are streaming. This finding implies that listeners are capable of extracting the tempo of the total alternating sequence, even though successive tones belong to different streams. The present study investigated whether subjects are also capable of establishing the tempo of one stream. To this end, listeners were required to match the rate of one stream by adjusting the rate of a nonalternating comparison stream (a pulse sequence). In addition, they were required to match the total rate of the streaming sequence. It turned out that the total tempo could be matched precisely and that the tempo of one stream could be approximated very closely.     

Effect on numerosity judgment of grouping of tones by auditory channels

Previous research suggests that numerosity judgments for sequences of tones improve when the sequence is structured such that equal and small groups of tones alternate between the ears. The present. study systematically investigates the effects of the structure of tone sequences on number judgment. Tone frequency is chosen as the grouping principle instead of spatial location. In the first experiment, sequences with equal groups of 2, 3, 4, 5, 6, or 7 tones (groups alternating between 800 and 1, 250 Hz) were compared with monotonous sequences. At a slow repetition rate of the tones (280 msec onset to onset), grouping the sequence deteriorated the numerosity judgment. At II fast repetition rate of the tones (100 msec onset to onset) grouping improved numerosity judgment, but only if the group size did not exceed 4 to 5 tones. In the second experiment, the equality of group size in the sequence as a necessary condition for the improvement was investigated. It was found that tone sequences comprised of equal groups were judged more accurately in number than sequences comprised of unequal groups. These results seem to give support for the existence of an auditory subitizing process. The results of this study are also compared with the results of studies in which sequences of tones, alternating one by om! between locations or frequencies, had to be judged in number.     

Pattern specificity in the effect of prior Δƒ on auditory stream segregation

During repeating sequences of low (A) and high (B) tones, perception of two separate streams ("streaming") increases with greater frequency separation (Δ?) between the A and B tones; in contrast, a prior context with large Δ? results in less streaming during a subsequent test pattern. The purpose of the present study was to investigate what aspects of the context pattern are necessary for this context effect to occur. Simply changing the B-tone frequency without an alternating A tone present was not sufficient to cause the effect of prior Δ?, but rather a melodic change between A and B tones was necessary. We further investigated the extent to which the context and test patterns needed to have similar rhythms (xxx-xxx-) and melodies (up-down-flat-up-down), and found that a maximal prior-Δ? effect occurred when the rhythmic patterns of the context and test were similar, regardless of the melodic structure. Thus, the effect of prior Δ? on streaming depended on the presence of (1) at least one melodic change in the context, and (2) similar rhythmic patterns in the context and test.     

Perceptual grouping in audition

The present experiments evaluated the effect of relative frequency as a determinant of the figure-ground organization of sequences of auditory tones. Observers counted sequences of 20 ms tones that were presented at the same frequency or that alternated between two different frequencies. The alternating tones differed in frequency by one whole tone, seven tones, or nineteen tones. Counting accuracy increased with increases in the silent interval between the tones. When the alternating tones differed by seven or nineteen tones, counting was disrupted at rates of presentation of eight tones per second or slower. In contrast to this decrement in the counting of tones that alternated by over an octave, very little decrement was observed when the tones alternated by just one whole tone. The best subjects counted these alternating tones more accurately than the tones presented at the same frequency. The poorest subjects showed a small decrement even when the tones alternated by just one whole tone. The results were discussed in terms of determinants of figure-ground organization in auditory information processing.     

Effects of attention and unilateral neglect on auditory stream segregation

Two pairs of experiments studied the effects of attention and of unilateral neglect on auditory streaming. The first pair showed that the build up of auditory streaming in normal participants is greatly reduced or absent when they attend to a competing task in the contralateral ear. It was concluded that the effective build up of streaming depends on attention. The second pair showed that patients with an attentional deficit toward the left side of space (unilateral neglect) show less stream segregation of tone sequences presented to their left than to their right ears. Streaming in their right ears was similar to that for stimuli presented to either ear of healthy and of brain-damaged controls, who showed no across-ear asymmetry. This result is consistent with an effect of attention on streaming, constrains the neural sites involved, and reveals a qualitative difference between the perception of left- and right-sided sounds by neglect patients.     

Attentional modulation in perception of visual motion events.

Identical visual targets moving across each other with equal and constant speed can be perceived either to bounce off or to stream through each other. This bistable motion perception has been studied mostly in the context of motion integration. Since the perception of most ambiguous motion is affected by attention, there is the possibility of attentional modulation occurring in this case as well. We investigated whether distraction of attention from the moving targets would alter the relative frequency of each percept. During the observation of the streaming/bouncing motion event in the peripheral visual field, visual attention was disrupted by an abrupt presentation of a visual distractor at various timings and locations (experiment 1; exogenous distraction of attention) or by the demand of an additional discrimination task (experiments 2 and 3; endogenous distraction of attention). Both types of distractions of attention increased the frequency of the bouncing percept and decreased that of the streaming percept. These results suggest that attention may facilitate the perception of object motion as continuing in the same direction as in the past.     

Responsiveness of Western adults to pitch-distributional information in melodic sequences

Responsiveness of musically trained and untrained adults to pitch-distributional information in melodic contexts was assessed. In Experiment 1, melodic contexts were pure-tone sequences, generated from either a diatonic or one of four nondiatonic tonesets, in which pitch-distributional information was manipulated by variation of the relative frequency of occurrence of tones from the toneset. Both the assignment of relative frequency of occurrence to tones and the construction of the (fixed) temporal order of tones within the sequences contravened the conventions of western tonal music. A probe-tone technique was employed. Each presentation of a sequence was followed by a probe tone, one of the 12 chromatic notes within the octave. Listeners rated the goodness of musical fit of the probe tone to the sequence. Probe-tone ratings were significantly related to frequency of occurrence of the probe tone in the sequence for both trained and untrained listeners. In addition, probe-tone ratings decreased as the pitch distance between the probe tone and the final tone of the sequence increased. For musically trained listeners, probe-tone ratings for diatonic sequences tended also to reflect the influence of an internalized tonal schema. Experiment 2 demonstrated that the temporal location of tones in the sequences could not alone account for the effect of frequency of occurrence in Experiment 1. Experiment 3 tested musically untrained listeners under the conditions of Experiment 1, with the exception that the temporal order of tones in each sequence was randomized across trials. The effect of frequency of occurrence found in Experiment 1 was replicated and strengthened.