Predictive cues for auditory stream formation in humans and monkeys.

Auditory perception is improved when stimuli are predictable, and this effect is evident in a modulation of the activity of neurons in the auditory cortex as shown previously. Human listeners can better predict the presence of duration deviants embedded in stimulus streams with fixed interonset interval (isochrony) and repeated duration pattern (regularity), and neurons in the auditory cortex of macaque monkeys have stronger sustained responses in the 60-140 ms post-stimulus time window under these conditions. Subsequently, the question has arisen whether isochrony or regularity in the sensory input contributed to the enhancement of the neuronal and behavioural responses. Therefore, we varied the two factors isochrony and regularity independently and measured the ability of human subjects to detect deviants embedded in these sequences as well as measuring the responses of neurons the primary auditory cortex of macaque monkeys during presentations of the sequences. The performance of humans in detecting deviants was significantly increased by regularity. Isochrony enhanced detection only in the presence of the regularity cue. In monkeys, regularity increased the sustained component of neuronal tone responses in auditory cortex while isochrony had no consistent effect. Although both regularity and isochrony can be considered as parameters that would make a sequence of sounds more predictable, our results from the human and monkey experiments converge in that regularity has a greater influence on behavioural performance and neuronal responses.

Probing neural mechanisms underlying auditory stream segregation in humans by transcranial direct current stimulation (tDCS).

One hypothesis concerning the neural underpinnings of auditory streaming states that frequency tuning of tonotopically organized neurons in primary auditory fields in combination with physiological forward suppression is necessary for the separation of representations of high-frequency A and low-frequency B tones. The extent of spatial overlap between the tonotopic activations of A and B tones is thought to underlie the perceptual organization of streaming sequences into one coherent or two separate streams. The present study attempts to interfere with these mechanisms by transcranial direct current stimulation (tDCS) and to probe behavioral outcomes reflecting the perception of ABAB streaming sequences. We hypothesized that tDCS by modulating cortical excitability causes a change in the separateness of the representations of A and B tones, which leads to a change in the proportions of one-stream and two-stream percepts. To test this, 22 subjects were presented with ambiguous ABAB sequences of three different frequency separations (∆F) and had to decide on their current percept after receiving sham, anodal, or cathodal tDCS over the left auditory cortex. We could confirm our hypothesis at the most ambiguous ∆F condition of 6 semitones. For anodal compared with sham and cathodal stimulation, we found a significant decrease in the proportion of two-stream perception and an increase in the proportion of one-stream perception. The results demonstrate the feasibility of using tDCS to probe mechanisms underlying auditory streaming through the use of various behavioral measures. Moreover, this approach allows one to probe the functions of auditory regions and their interactions with other processing stages.

Decision making and ambiguity in auditory stream segregation.

Researchers of auditory stream segregation have largely taken a bottom-up view on the link between physical stimulus parameters and the perceptual organization of sequences of ABAB sounds. However, in the majority of studies, researchers have relied on the reported decisions of the subjects regarding which of the predefined percepts (e.g., one stream or two streams) predominated when subjects listened to more or less ambiguous streaming sequences. When searching for neural mechanisms of stream segregation, it should be kept in mind that such decision processes may contribute to brain activation, as also suggested by recent human imaging data. The present study proposes that the uncertainty of a subject in making a decision about the perceptual organization of ambiguous streaming sequences may be reflected in the time required to make an initial decision. To this end, subjects had to decide on their current percept while listening to ABAB auditory streaming sequences. Each sequence had a duration of 30 s and was composed of A and B harmonic tone complexes differing in fundamental frequency (ΔF). Sequences with seven different ΔF were tested. We found that the initial decision time varied non-monotonically with ΔF and that it was significantly correlated with the degree of perceptual ambiguity defined from the proportions of time the subjects reported a one-stream or a two-stream percept subsequent to the first decision. This strong relation of the proposed measures of decision uncertainty and perceptual ambiguity should be taken into account when searching for neural correlates of auditory stream segregation.

Corrigendum: Probing auditory scene analysis.

[This corrects the article on p. 293 in vol. 8, PMID: 25309314.].

Time course of auditory streaming: do CI users differ from normal-hearing listeners?

In a complex acoustical environment, the auditory system decides which stimulus components originate from the same source by forming auditory streams, where temporally non-overlapping stimulus portions are considered to originate from one source if their stimulus characteristics are similar. The mechanisms underlying streaming are commonly studied by alternating sequences of A and B signals which are often tones with different frequencies. For similar frequencies, they are grouped into one stream. Otherwise, they are considered to belong to different streams. The present study investigates streaming in cochlear implant (CI) users, where hearing is restored by electrical stimulation of the auditory nerve. CI users listened to 30-s long sequences of alternating A and B harmonic complexes at four different fundamental frequency separations, ranging from 2 to 14 semitones. They had to indicate as promptly as possible after sequence onset, if they perceived one stream or two streams and, in addition, any changes of the percept throughout the rest of the sequence. The conventional view is that the initial percept is always that of a single stream which may after some time change to a percept of two streams. This general build-up hypothesis has recently been challenged on the basis of a new analysis of data of normal-hearing listeners. Using the same experimental paradigm and analysis, the present study found that the results of CI users agree with those of the normal-hearing listeners: (i) the probability of the first decision to be a one-stream percept decreased and that of a two-stream percept increased as Δf increased, and (ii) a build-up was only found for 6 semitones. Only the time elapsed before the listeners made their first decision of the percept was prolonged as compared to normal-hearing listeners. The similarity in the data of the CI user and the normal-hearing listeners indicates that the quality of stream formation is similar in these groups of listeners.

Evaluating auditory stream segregation of SAM tone sequences by subjective and objective psychoacoustical tasks, and brain activity.

Auditory stream segregation refers to a segregated percept of signal streams with different acoustic features. Different approaches have been pursued in studies of stream segregation. In psychoacoustics, stream segregation has mostly been investigated with a subjective task asking the subjects to report their percept. Few studies have applied an objective task in which stream segregation is evaluated indirectly by determining thresholds for a percept that depends on whether auditory streams are segregated or not. Furthermore, both perceptual measures and physiological measures of brain activity have been employed but only little is known about their relation. How the results from different tasks and measures are related is evaluated in the present study using examples relying on the ABA- stimulation paradigm that apply the same stimuli. We presented A and B signals that were sinusoidally amplitude modulated (SAM) tones providing purely temporal, spectral or both types of cues to evaluate perceptual stream segregation and its physiological correlate. Which types of cues are most prominent was determined by the choice of carrier and modulation frequencies (f mod) of the signals. In the subjective task subjects reported their percept and in the objective task we measured their sensitivity for detecting time-shifts of B signals in an ABA- sequence. As a further measure of processes underlying stream segregation we employed functional magnetic resonance imaging (fMRI). SAM tone parameters were chosen to evoke an integrated (1-stream), a segregated (2-stream), or an ambiguous percept by adjusting the f mod difference between A and B tones (Δf mod). The results of both psychoacoustical tasks are significantly correlated. BOLD responses in fMRI depend on Δf mod between A and B SAM tones. The effect of Δf mod, however, differs between auditory cortex and frontal regions suggesting differences in representation related to the degree of perceptual ambiguity of the sequences.

Rhythm sensitivity in macaque monkeys.

This study provides evidence that monkeys are rhythm sensitive. We composed isochronous tone sequences consisting of repeating triplets of two short tones and one long tone which humans perceive as repeating triplets of two weak and one strong beat. This regular sequence was compared to an irregular sequence with the same number of randomly arranged short and long tones with no such beat structure. To search for indication of rhythm sensitivity we employed an oddball paradigm in which occasional duration deviants were introduced in the sequences. In a pilot study on humans we showed that subjects more easily detected these deviants when they occurred in a regular sequence. In the monkeys we searched for spontaneous behaviors the animals executed concomitant with the deviants. We found that monkeys more frequently exhibited changes of gaze and facial expressions to the deviants when they occurred in the regular sequence compared to the irregular sequence. In addition we recorded neuronal firing and local field potentials from 175 sites of the primary auditory cortex during sequence presentation. We found that both types of neuronal signals differentiated regular from irregular sequences. Both signals were stronger in regular sequences and occurred after the onset of the long tones, i.e., at the position of the strong beat. Local field potential responses were also significantly larger for the durational deviants in regular sequences, yet in a later time window. We speculate that these temporal pattern-selective mechanisms with a focus on strong beats and their deviants underlie the perception of rhythm in the chosen sequences.

The Build-up of Auditory Stream Segregation: A Different Perspective.

The build-up of auditory stream segregation refers to the notion that sequences of alternating A and B sounds initially tend to be heard as a single stream, but with time appear to split into separate streams. The central assumption in the analysis of this phenomenon is that streaming sequences are perceived as one stream at the beginning by default. In the present study, we test the validity of this assumption and document its impact on the apparent build-up phenomenon. Human listeners were presented with ABAB sequences, where A and B were harmonic tone complexes of seven different fundamental frequency separations (Δf) ranging from 2 to 14 semitones. Subjects had to indicate, as promptly as possible, their initial percept of the sequences, as either "one stream" or "two streams," and any changes thereof during the sequences. We found that subjects did not generally indicate a one-stream percept at the beginning of streaming sequences. Instead, the first perceptual decision depended on Δf, with the probability of a one-stream percept decreasing, and that of a two-stream percept increasing, with increasing Δf. Furthermore, subjects required some time to make and report a decision on their perceptual organization. Taking this time into account, the resulting time courses of two-stream probabilities differ markedly from those suggested by the conventional analysis. A build-up-like increase in two-stream probability was found only for the Δf of six semitones. At the other Δf conditions no or only minor increases in two-stream probability occurred. These results shed new light on the build-up of stream segregation and its possible neural correlates.

Active stream segregation specifically involves the left human auditory cortex.

An important aspect of auditory scene analysis is the sequential grouping of similar sounds into one "auditory stream" while keeping competing streams separate. In the present low-noise fMRI study we presented sequences of alternating high-pitch (A) and low-pitch (B) complex harmonic tones using acoustic parameters that allow the perception of either two separate streams or one alternating stream. However, the subjects were instructed to actively and continuously segregate the A from the B stream. This was controlled by the additional instruction to listen for rare level deviants only in the low-pitch stream. Compared to the control condition in which only one non-separable stream was presented the active segregation of the A from the B stream led to a selective increase of activation in the left auditory cortex (AC). Together with a similar finding from a previous study using a different acoustic cue for streaming, namely timbre, this suggests that the left auditory cortex plays a dominant role in active sequential stream segregation. However, we found cue differences within the left AC: Whereas in the posterior areas, including the planum temporale, activation increased for both acoustic cues, the anterior areas, including Heschl's gyrus, are only involved in stream segregation based on pitch.

A multilevel and cross-modal approach towards neuronal mechanisms of auditory streaming.

We report first results of a multilevel, cross-modal study on the neuronal mechanisms underlying auditory sequential streaming, with the focus on the impact of visual sequences on perceptually ambiguous tone sequences which can either be perceived as two separate streams or one alternating stream. We combined two psychophysical experiments performed on humans and monkeys with two human brain imaging experiments which allow to obtain complementary information on brain activation with high spatial (fMRI) and high temporal (MEG) resolution. The same acoustic paradigm based on the pairing of tone sequences with visual stimuli was used in all human studies and, in an adapted version, in the psychophysical study on monkeys. Our multilevel approach provides experimental evidence that the pairing of auditory and visual stimuli can reliably introduce a bias towards either an integrated or a segregated perception of ambiguous sequences. Thus, comparable to an explicit instruction, this approach can be used to control the subject's perceptual organization of an ambiguous sound sequence without the need for the subject to directly report it. This finding is of particular importance for animal studies because it allows to compare electrophysiological responses of auditory cortex neurons to the same acoustic stimulus sequence eliciting either a segregated or integrated percept.

Auditory stream segregation relying on timbre involves left auditory cortex.

An important aspect of auditory scene analysis is sequential grouping of sounds that are similar to one another in preference to sounds that follow one another. This grouping problem is captured by stream segregation tasks with alternating distinct sounds. We examined human auditory cortex activity with low noise fMRI in a stream segregation experiment relying on timbre differences of alternating harmonic tones (organ-like and trumpet-like). We found that stream segregation performance in comparison to monitoring a non-separable control stream increased activation exclusively in left auditory cortex and particularly in posterior areas. Our results suggest that left auditory cortex is selectively involved in this complex sequential task although the available cue for sequential grouping was timbre, usually attributed to right hemisphere analysis.