Pattern specificity in the effect of prior Δ? 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. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

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 (Δf). Larger Δf 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 Δf 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 Δf 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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Behavioral measures of auditory streaming in ferrets (Mustela putorius).

An important aspect of the analysis of auditory “scenes” relates to the perceptual organization of sound sequences into auditory “streams.” In this study, we adapted two auditory perception tasks, used in recent human psychophysical studies, to obtain behavioral measures of auditory streaming in ferrets (Mustela putorius). One task involved the detection of shifts in the frequency of tones within an alternating tone sequence. The other task involved the detection of a stream of regularly repeating target tones embedded within a randomly varying multitone background. In both tasks, performance was measured as a function of various stimulus parameters, which previous psychophysical studies in humans have shown to influence auditory streaming. Ferret performance in the two tasks was found to vary as a function of these parameters in a way that is qualitatively consistent with the human data. These results suggest that auditory streaming occurs in ferrets, and that the two tasks described here may provide a valuable tool in future behavioral and neurophysiological studies of the phenomenon. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

An objective measurement of the build-up of auditory streaming and of its modulation by attention.

Three experiments studied auditory streaming using sequences of alternating “ABA” triplets, where “A” and “B” were 50-ms tones differing in frequency by Δf semitones and separated by 75-ms gaps. Experiment 1 showed that detection of a short increase in the gap between a B tone and the preceding A tone, imposed on one ABA triplet, was better when the delay occurred early versus late in the sequence, and for Δf = 4 vs. Δf = 8. The results of this experiment were consistent with those of a subjective streaming judgment task. Experiment 2 showed that the detection of a delay 12.5 s into a 13.5-s sequence could be improved by requiring participants to perform a task on competing stimuli presented to the other ear for the first 10 s of that sequence. Hence, adding an additional task demand could improve performance via its effect on the perceptual organization of a sound sequence. The results demonstrate that attention affects streaming in an objective task and that the effects of build-up are not completely under voluntary control. In particular, even though build-up can impair performance in an objective task, participants are unable to prevent this from happening. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

"Recalibrating" the auditory system: The perception of loudness.

Listening to relatively intense tones at one frequency and weak tones at another makes the latter relatively louder. The auditory system's relative response to low-frequency (f?) and high-frequency (f?) tones depends on the separation between f? and f?. When f? and f? differ little, loudness matches change little with shifts in mean sound pressure levels (SPLs) at each frequency; but when f? and f? differ more, matches change markedly, showing how the auditory system "recalibrates" its responses to f? and f?. The magnitude of recalibration and its frequency bandwidth also depend to a modest degree on the range of SPLs, their mean level, and the experimental paradigm. The representation of loudness reflects the processing and recalibration of multidimensional peripheral inputs within a higher level, context-sensitive (adaptation-like) mechanism. Other perceptual modalities show evidence of analogous mechanisms. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Auditory stream segregation and the perception of across-frequency synchrony.

This study explored the extent to which sequential auditory grouping affects the perception of temporal synchrony. In Experiment 1, listeners discriminated between 2 pairs of asynchronous “target” tones at different frequencies, A and B, in which the B tone either led or lagged. Thresholds were markedly higher when the target tones were temporally surrounded by “captor tones” at the A frequency than when the captor tones were absent or at a remote frequency. Experiment 2 extended these findings to asynchrony detection, revealing that the perception of synchrony, one of the most potent cues for simultaneous auditory grouping, is not immune to competing effects of sequential grouping. Experiment 3 examined the influence of ear separation on the interactions between sequential and simultaneous grouping cues. The results showed that, although ear separation could facilitate perceptual segregation and impair asynchrony detection, it did not prevent the perceptual integration of simultaneous sounds. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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.     

Correction to Roberts et al. (2008).

Reports an error in "Effects of the build-up and resetting of auditory stream segregation on temporal discrimination" by Brian Roberts, Brian R. Glasberg and Brian C. J. Moore (Journal of Experimental Psychology: Human Perception and Performance, 2008[Aug], Vol 34[4], 992-1006). The year listed is incorrect. The article should have been dated 2008. (The following abstract of the original article appeared in record 2008-09670-016.) 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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Mechanisms of signal analysis and pattern perception in periodicity pitch

Progress in the knowledge of auditory processing of complex sounds has been made through coordinated psychophysical, physiological and theoretical studies of periodicity pitch and combination tones. Periodicity pitch is the basis for human perception of musical notes and pitch of voiced speech. The mechanism of perception involves harmonic pattern recognition on the complex Fourier frequency spectra generated by auditory frequency analysis. Combination tones are perceptible distortion tones generated within the cochlea by nonlinear interaction of component stimulus tones. Perception of periodicity pitch is quantitatively accounted for by a two-stage process of frequency analysis subject to random errors and significant nonlinearities, followed by a pattern recognizer that operates very efficiently to measure the period of musical and speech sounds. The basic characteristic of the first stage is a Gaussian standard error function that quantifies the randomness in aural estimation of frequencies of component tones in a complex tone stimulus. Efficient aural measurement of neural spike intervals from the eighth nerve provides a physiological account for the psychophysical characteristic of aural frequency analysis with complex sounds. Although cochlear filtering is an essential stage in auditory frequency analysis, neural time following, rather than details of the filter characteristics, is the decisive factor in determining the precision of aural frequency measurement. It is likely that peripheral auditory coding is similar for sounds in periodicity pitch and in speech perception, although the 'second stage' representing central processing would differ.     

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. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Perceptual grouping of tone sequences by normally hearing and hearing-impaired listeners

This study examined the perceptual grouping of rapid tone sequences for listeners with normal hearing and listeners with unilateral and bilateral cochlear hearing loss. The sequence ABA-ABA- was used, where A and B represent sinusoidal tones bursts (10-ms rise/fall, 80-ms steady state, 20-ms interval between tones) and - represents a silent interval of 120 ms. Tone A was fixed in frequency at 250, 500, 1000, or 2000 Hz. Tone B started with a frequency well above or below that of tone A, and its frequency was swept towards that of tone A so that the frequency separation between them decreased in an exponential manner. Listeners were required to indicate when they could no longer hear the tones A and B as two separate streams, but heard only a single stream with a "gallop" rhythm. This is called the fission boundary. For the normally hearing listeners, the separation between tones A and B at the fission boundary was roughly independent of the frequency of tone A when expressed as the difference in number of ERBs (delta E) between A and B, which is consistent with a recent model of stream segregation [M. W. Beauvois and R. Meddis, J. Acoust. Soc. Am. 99, 2270-2280 (1996)]. For the unilaterally hearing-impaired listeners, there was no consistent difference in the delta E magnitudes across ears, even though the auditory filters were broader in the impaired ears. This is not consistent with the theory of Beauvois and Meddis. The bilaterally hearing-impaired listeners sometimes showed delta E magnitudes within the normal range, and sometimes showed larger than normal delta E magnitudes. The results are discussed in terms of the factors that might influence perceptual stream formation in hearing-impaired listeners.     

Sequential stream segregation in the absence of spectral cues

This paper investigates the cues used by the auditory system in the perceptual organization of sequential sounds. In particular, the ability to organize sounds in the absence of spectral cues is studied. In the first experiment listeners were presented with a tone sequence ABA ABA ..., where the fundamental frequency (f0) of tone A was fixed at 100 Hz and the f0 difference between tones A and B varied across trials between 1 and 11 semitones. Three spectral conditions were tested: pure tones, harmonic complexes filtered with a bandpass region between 500 and 2000 Hz, and harmonic complexes filtered with a bandpass region chosen so that only harmonics above the tenth would be passed by the filter, thus severely limiting spectral information. Listeners generally reported that they could segregate tones A and B into two separate perceptual streams when the f0 interval exceeded about four semitones. This was true for all conditions. The second experiment showed that most listeners were better able to recognize a short atonal melody interleaved with random distracting tones when the distracting tones were in an f0 region 11 semitones higher than the melody than when the distracting tones were in the same f0 region. The results were similar for both pure tones and complex tones comprising only high, unresolved harmonics. The results from both experiments show that spectral separation is not a necessary condition for perceptual stream segregation. This suggests that models of stream segregation that are based solely on spectral properties may require some revision.     

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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Representation of spectral and temporal sound features in three cortical fields of the cat. Similarities outweigh differences

This study investigates the degree of similarity of three different auditory cortical areas with respect to the coding of periodic stimuli. Simultaneous single- and multiunit recordings in response to periodic stimuli were made from primary auditory cortex (AI), anterior auditory field (AAF), and secondary auditory cortex (AII) in the cat to addresses the following questions: is there, within each cortical area, a difference in the temporal coding of periodic click trains, amplitude-modulated (AM) noise bursts, and AM tone bursts? Is there a difference in this coding between the three cortical fields? Is the coding based on the temporal modulation transfer function (tMTF) and on the all-order interspike-interval (ISI) histogram the same? Is the perceptual distinction between rhythm and roughness for AM stimuli related to a temporal versus spatial representation of AM frequency in auditory cortex? Are interarea differences in temporal response properties related to differences in frequency tuning? The results showed that: 1) AM stimuli produce much higher best modulation frequencies (BMFs) and limiting rates than periodic click trains. 2) For periodic click trains and AM noise, the BMFs and limiting rates were not significantly different for the three areas. However, for AM tones the BMF and limiting rates were about a factor 2 lower in AAF compared with the other areas. 3) The representation of stimulus periodicity in ISIs resulted in significantly lower mean BMFs and limiting rates compared with those estimated from the tMTFs. The difference was relatively small for periodic click trains but quite large for both AM stimuli, especially in AI and AII. 4) Modulation frequencies <20 Hz were represented in the ISIs, suggesting that rhythm is coded in auditory cortex in temporal fashion. 5) In general only a modest interdependence of spectral- and temporal-response properties in AI and AII was found. The BMFs were correlated positively with characteristic frequency in AAF. The limiting rate was positively correlated with the frequency-tuning curve bandwidth in AI and AII but not in AAF. Only in AAF was a correlation between BMF and minimum latency was found. Thus whereas differences were found in the frequency-tuning curve bandwidth and minimum response latencies among the three areas, the coding of periodic stimuli in these areas was fairly similar with the exception of the very poor representation of AM tones in AII. This suggests a strong parallel processing organization in auditory cortex.     

Mismatch negativity: deviance detection or the maintenance of the 'standard'

Electric brain responses were measured to infrequent tones that broke the frequency alternation of two tones, deviated in duration or violated both regularities (alternation and constant duration). Mismatch negativity (MMN) was elicited by both simple deviants with the duration-related MMN peaking approximately 130 ms later than the alternation-related MMN. The double deviant elicited two successive MMNs. Thus violation of each regularity elicited a separate MMN, whereas previous studies showed that multiple temporally separate deviations from a single repetitive standard elicit one MMN only. These results suggest that the primary function of the MMN-generating process is more closely related to maintaining the representation of auditory regularities than to deviance detection per se.     

Tonotopic organization of auditory cortical fields delineated by parvalbumin immunoreactivity in macaque monkeys

Tonotopic maps, obtained from single and multi-unit recordings in the primary and surrounding areas of the auditory cortex, were related to chemoarchitecture of the supratemporal plane, as delineated by immunoreactivity for parvalbumin. Neurons in the central core were sharply tuned and formed two complete tonotopic representations corresponding to the primary auditory area (AI) and the rostral (R) area. High frequencies were represented posteriorly in AI and anteriorly in R, the representation reversing in the anterior part of the core. Neurons in regions of less dense immunostaining previously described as lateral (L) and posteromedial (P-m) fields, showed broader frequency tuning. Two tonotopic representations were found in L: in an anterolateral (AL) field, corresponding to a field previously reported by others, high frequencies were represented anteriorly and low frequencies posteriorly; in a posterolateral field (PL) the trend reversed. There was a further reversal on entering P-m from the high frequency representation in PL and progressively lower frequencies tended to be represented more medially in P-m, but P-m may contain two representations reported by others. Neurons in the previously described anteromedial (A-m) and medial (M) fields of weaker immunostaining, were even more broadly tuned. A tonotopic progression from low frequency representation posteriorly to high frequency representation anteriorly was observed in the medial field. Frequency representation in A-m remains uncertain. No tonotopic representation could be demonstrated with the stimuli used in the zones of very weak parvalbumin immunostaining outside AL, PL, P-m, A-m, and M. The properties of neurons in the core and surrounding zones are likely to reflect inputs from the ventral and dorsal medial geniculate nuclei, respectively. The fields outside the core seem to be the starting points for separate streams of auditory corticocortical connections passing into association cortex.     

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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Dual-task interference and visual encoding.

A visual target (T?) containing either 1 or 2 letters, or a random 10-sided polygon, was presented after an auditory target (T?) at a stimulus onset asynchrony (SOA) of either 50, 150, 250, or 600 ms. Task? was a speeded pitch discrimination to the tone, and across experiments, T? was either 1 of 2 tones (2-alternative discrimination [2AD]) or 1 of 4 tones (4-alternative discrimination [4AD]). Memory for the visual information decreased as SOA was reduced when a mask was used, but not when there was no mask. The effects of SOA were larger for the 4AD Task? than the 2AD Task?. The results demonstrate cross-modal, dual-task interference on visual encoding and suggest central interference with the short-term consolidation of visual information in short-term memory. (PsycINFO Database Record (c) 2010 APA, all rights reserved)