Search

Visual search is widely studied, with paradigms ranging from simple stimuli and tasks to complex scenes and real-world settings. Here, we investigated a real-world scenario, the search for fire sources with a novel augmented reality (AR) device: a firefighter’s helmet equipped with a heat sensor and an integrated display indicating the heat distribution in the sensor’s field of view. This was compared to the current state-of-the-art, a hand-held thermal imaging camera. Although both are actual devices, we used a high fidelity virtual-reality (VR) simulation to provide a safe and controlled, yet realistic, experimental setting. Detecting fire sources was faster with the thermal camera than with the helmet display. Responses in target-present trials were faster than in target-absent trials for both devices. Fire localization after detection was numerically faster and more accurate, in particular in the horizontal plane, for the helmet display than for the thermal camera. Search was strongly biased to start on the left-hand side of each room, reminiscent of pseudoneglect and similar biases in scene viewing. Our study exemplifies how VR can be used to study vision in realistic settings, to foster the development of AR devices, and to obtain results relevant to basic science and applications alike.

In multistability, perceptual interpretations (“percepts”) of ambiguous stimuli alternate over time. There is considerable debate as to whether similar regularities govern the first percept after stimulus onset and percepts during prolonged presentation. We address this question in a visual pattern-component rivalry paradigm by presenting two overlaid drifting gratings, which participants perceived as individual gratings passing in front of each other (“segregated”) or as a plaid (“integrated”). We varied the enclosed angle (“opening angle”) between the gratings (experiments 1 and 2) and stimulus orientation (experiment 2). The relative number of integrated percepts increased monotonically with opening angle. The point of equality, where half of the percepts were integrated, was at a smaller opening angle at onset than during prolonged viewing. The functional dependence of the relative number of integrated percepts on opening angle showed a steeper curve at onset than during prolonged viewing. Dominance durations of integrated percepts were longer at onset than during prolonged viewing and increased with opening angle. The general pattern persisted when stimuli were rotated (experiment 2), despite some perceptual preference for cardinal motion directions over oblique directions. Analysis of eye movements, specifically the slow phase of the optokinetic nystagmus (OKN), confirmed the veridicality of participants’ reports and provided a temporal characterization of percept formation after stimulus onset. Together, our results show that the first percept after stimulus onset exhibits a different dependence on stimulus parameters than percepts during prolonged viewing. This underlines the distinct role of the first percept in multistability.

Human brain function draws on predictive mechanisms that exploit higher-level context during lower-level perception. These mechanisms are particularly relevant for situations in which sensory information is compromised or incomplete, as for example in natural speech where speech segments may be omitted due to sluggish articulation. Here, we investigate which brain areas support the processing of incomplete words that were predictable from semantic context, compared with incomplete words that were unpredictable. During functional magnetic resonance imaging (fMRI), participants heard sentences that orthogonally varied in predictability (semantically predictable vs. unpredictable) and completeness (complete vs. incomplete, i.e. missing their final consonant cluster). The effects of predictability and completeness interacted in heteromodal semantic processing areas, including left angular gyrus and left precuneus, where activity did not differ between complete and incomplete words when they were predictable. The same regions showed stronger activity for incomplete than for complete words when they were unpredictable. The interaction pattern suggests that for highly predictable words, the speech signal does not need to be complete for neural processing in semantic processing areas. Hum Brain Mapp 37:704-716, 2016. © 2015 Wiley Periodicals, Inc.

By encoding acoustic regularities present in the environment, the human brain can generate predictions of what is likely to occur next. Recent studies suggest that deviations from encoded regularities are detected within 10–50 ms after stimulus onset, as indicated by electrophysiological effects in the middle latency response (MLR) range. This is upstream of previously known long-latency (LLR) signatures of deviance detection such as the mismatch negativity (MMN) component. In the present study, we created predictable and unpredictable contexts to investigate MLR and LLR signatures of the encoding of spatial auditory regularities and the generation of predictions from these regularities. Chirps were monaurally delivered in an either regular (predictable: left–right–left–right) or a random (unpredictable left/right alternation or repetition) manner. Occasional stimulus omissions occurred in both types of sequences. Results showed that the Na component (peaking at 34 ms after stimulus onset) was attenuated for regular relative to random chirps, albeit no differences were observed for stimulus omission responses in the same latency range. In the LLR range, larger chirp-and omission-evoked responses were elicited for the regular than for the random condition, and predictability effects were more prominent over the right hemisphere. We discuss our findings in the framework of a hierarchical organization of spatial regularity encoding. This article is part of a Special Issue entitled SI: Prediction and Attention.

Many sound sources emit series of discrete sounds. Auditory perception must bind these sounds together (stream integration) while separating them from sounds emitted by other sources (stream segregation). One cue for identifying successive sounds that belong together is the predictability between their feature values. Previous studies have demonstrated that independent predictable patterns appearing separately in two interleaved sound sequences support perceptual segregation. The converse case, whether a joint predictable pattern in a mixture of interleaved sequences supports perceptual integration, has not yet been put to a rigorous empirical test. This was mainly due to difficulties in manipulating the predictability of the full sequence independently of the predictability of the interleaved subsequences. The present study implemented such an independent manipulation. Listeners continuously indicated whether they perceived a tone sequence as integrated or segregated, while predictable patterns set up to support one or the other percept were manipulated without the participants’ knowledge. Perceptual reports demonstrate that predictability supports stream segregation or integration depending on the type of predictable pattern that is present in the sequence. The effects of predictability were so pronounced as to qualitatively flip perception from predominantly (62%) integrated to predominantly (73%) segregated. These results suggest that auditory perception flexibly responds to encountered regular patterns, favoring predictable perceptual organizations over unpredictable ones. Besides underlining the role of predictability as a cue within auditory scene analysis, the present design also provides a general framework that accommodates previous investigations focusing on sub-comparisons within the present set of experimental manipulations. Results of intermediate conditions shed light on why some previous studies have obtained little to no effects of predictability on auditory scene analysis.

For studying multistable auditory perception, we propose a paradigm that evokes integrated or segregated perception of a sound sequence, and permits decomposition of the segregated grouping into foreground and background sounds. The paradigm combines 3-tone pitch patterns with alternating timbres, resulting in a repeating 6-tone structure that can be perceived as rising based on temporal proximity, or as falling based on timbre similarity. Listeners continuously report their percept while EEG is recorded. Results show an ERP modulation starting at ∼70 ms after sound onset that can be explained by whether a sound belongs to perceived foreground or background, with no additional effect of integrated versus segregated grouping. Auditory grouping as indexed by the mismatch negativity did not correspond with reported sound grouping. The paradigm offers a new possibility for investigating effects of conscious perceptual organization on sound processing.

In our surrounding acoustic world sounds are produced by different sources and interfere with each other before arriving to the ears. A key function of the auditory system is to provide consistent and robust descriptions of the coherent sound groupings and sequences (auditory objects), which likely correspond to the various sound sources in the environment. This function has been termed auditory stream segregation. In the current study we tested the effects of separation in the frequency of amplitude modulation on the segregation of concurrent sound sequences in the auditory stream–segregation paradigm (van Noorden 1975). The aim of the study was to assess 1) whether differential amplitude modulation would help in separating concurrent sound sequences and 2) whether this cue would interact with previously studied static cues (carrier frequency and location difference) in segregating concurrent streams of sound. We found that amplitude modulation difference is utilized as a primary cue for the stream segre...

Sound sources often emit trains of discrete sounds, such as a series of footsteps. Previously, two different principles have been suggested for how the human auditory system binds discrete sounds together into perceptual units. The feature similarity principle is based on linking sounds with similar characteristics over time. The predictability principle is based on linking sounds that follow each other in a predictable manner. The present study compared the effects of these two principles. Participants were presented with tone sequences and instructed to continuously indicate whether they perceived a single coherent sequence or two concurrent streams of sound. We investigated the influence of separate manipulations of similarity and predictability on these perceptual reports. Both grouping principles affected perception of the tone sequences, albeit with different characteristics. In particular, results suggest that whereas predictability is only analyzed for the currently perceived sound organization, f...

The remarkable capabilities displayed by humans in making sense of an overwhelming amount of sensory information cannot be explained easily if perception is viewed as a passive process. Current theoretical and computational models assume that to achieve meaningful and coherent perception, the human brain must anticipate upcoming stimulation. But how are upcoming stimuli predicted in the brain? We unmasked the neural representation of a prediction by omitting the predicted sensory input. Electrophysiological brain signals showed that when a clear prediction can be formulated, the brain activates a template of its response to the predicted stimulus before it arrives to our senses.

Empirical research on the sequential decomposition of an auditory scene primarily relies on interleaved sound mixtures of only two tone sequences (e.g., ABAB…). This oversimplifies the sound decomposition problem by limiting the number of putative perceptual organizations. The current study used a sound mixture composed of three different tones (ABCABC…) that could be perceptually organized in many different ways. Participants listened to these sequences and reported their subjective perception by continuously choosing one out of 12 visually presented perceptual organization alternatives. Different levels of frequency and spatial separation were implemented to check whether participants' perceptual reports would be systematic and plausible. As hypothesized, while perception switched back and forth in each condition between various perceptual alternatives (multistability), spatial as well as frequency separation generally raised the proportion of segregated and reduced the proportion of integrated alternatives. During segregated percepts, in contrast to the hypothesis, many participants had a tendency to perceive two streams in the foreground, rather than reporting alternatives with a clear foreground-background differentiation. Finally, participants perceived the organization with intermediate feature values (e.g., middle tones of the pattern) segregated in the foreground slightly less often than similar alternatives with outer feature values (e.g., higher tones).

In binocular rivalry, paradigms have been proposed for unobtrusive moment-by-moment readout of observers' perceptual experience ("no-report paradigms"). Here, we take a first step to extend this concept to auditory multistability. Observers continuously reported which of two concurrent tone sequences they perceived in the foreground: high-pitch (1008 Hz) or low-pitch (400 Hz) tones. Interstimulus intervals were either fixed per sequence (Experiments 1 and 2) or random with tones alternating (Experiment 3). A horizontally drifting grating was presented to each eye; to induce binocular rivalry, gratings had distinct colors and motion directions. To associate each grating with one tone sequence, a pattern on the grating jumped vertically whenever the respective tone occurred. We found that the direction of the optokinetic nystagmus (OKN)-induced by the visually dominant grating-could be used to decode the tone (high/low) that was perceived in the foreground well above chance. This OKN-based readout improved after observers had gained experience with the auditory task (Experiments 1 and 2) and for simpler auditory tasks (Experiment 3). We found no evidence that the visual stimulus affected auditory multistability. Although decoding performance is still far from perfect, our paradigm may eventually provide a continuous estimate of the currently dominant percept in auditory multistability.

When distinct stimuli are presented to the two eyes, their mental representations alternate in awareness. Here, such "binocular rivalry" was used to investigate whether audio-visual associations bias visual perception. To induce two arbitrary associations, each between a tone and a grating of a specific color and motion direction, observers were required to respond whenever this combination was presented, but not for other tone-grating combinations. After about 20min of this induction phase, each of the gratings was presented to one eye to induce rivalry, while either of the two tones or no tone was played. Observers were asked to watch the rivaling stimuli and listen to the tones. The observer's dominant percept was assessed throughout by measuring the optokinetic nystagmus (OKN), whose slow phase follows the direction of the currently dominant grating. We found that perception in rivalry was affected by the concurrently played tone. Results suggest a bias towards the grating that had been associated with the concurrently presented tone and prolonged dominance durations for this grating compared to the other. Numerically, conditions without tone fell in-between for measures of bias and dominance duration. Our data show that a rapidly acquired arbitrary audio-visual association biases visual perception. Unlike previously reported cross-modal interactions in rivalry, this effect can neither be explained by a pure attentional (dual-task) effect, nor does it require a fixed physical or semantic relation between the auditory and visual stimulus. This suggests that audio-visual associations that are quickly formed by associative learning may affect visual representations directly.

While subjective reports provide a direct measure of perception, their validity is not self-evident. Here, the authors tested three possible biasing effects on perceptual reports in the auditory streaming paradigm: errors due to imperfect understanding of the instructions, voluntary perceptual biasing, and susceptibility to implicit expectations. (1) Analysis of the responses to catch trials separately promoting each of the possible percepts allowed the authors to exclude participants who likely have not fully understood the instructions. (2) Explicit biasing instructions led to markedly different behavior than the conventional neutral-instruction condition, suggesting that listeners did not voluntarily bias their perception in a systematic way under the neutral instructions. Comparison with a random response condition further supported this conclusion. (3) No significant relationship was found between social desirability, a scale-based measure of susceptibility to implicit social expectations, and any of the perceptual measures extracted from the subjective reports. This suggests that listeners did not significantly bias their perceptual reports due to possible implicit expectations present in the experimental context. In sum, these results suggest that valid perceptual data can be obtained from subjective reports in the auditory streaming paradigm.

Auditory stream segregation involves linking temporally separate acoustic events into one or more coherent sequences. For any non-trivial sequence of sounds, many alternative descriptions can be formed, only one or very few of which emerge in awareness at any time. Evidence from studies showing bi-/multistability in auditory streaming suggest that some, perhaps many of the alternative descriptions are represented in the brain in parallel and that they continuously vie for conscious perception. Here, based on a predictive coding view, we consider the nature of these sound representations and how they compete with each other. Predictive processing helps to maintain perceptual stability by signalling the continuation of previously established patterns as well as the emergence of new sound sources. It also provides a measure of how well each of the competing representations describes the current acoustic scene. This account of auditory stream segregation has been tested on perceptual data obtained in the auditory streaming paradigm.

The auditory system continuously parses the acoustic environment into auditory objects, usually representing separate sound sources. Sound sources typically show characteristic emission patterns. These regular temporal sound patterns are possible cues for distinguishing sound sources. The present study was designed to test whether regular patterns are used as cues for source distinction and to specify the role that detecting these regularities may play in the process of auditory stream segregation. Participants were presented with tone sequences, and they were asked to continuously indicate whether they perceived the tones in terms of a single coherent sequence of sounds (integrated) or as two concurrent sound streams (segregated). Unknown to the participant, in some stimulus conditions, regular patterns were present in one or both putative streams. In all stimulus conditions, participants' perception switched back and forth between the two sound organizations. Importantly, regular patterns occurring in either one or both streams prolonged the mean duration of two-stream percepts, whereas the duration of one-stream percepts was unaffected. These results suggest that temporal regularities are utilized in auditory scene analysis. It appears that the role of this cue lies in stabilizing streams once they have been formed on the basis of simpler acoustic cues.

The separation of concurrently presented sounds into distinct auditory objects is accompanied by the elicitation of the object-related negativity (ORN) component of the event-related potential (ERP). In the present study, participants were exposed to tone sequences containing not only concurrent (harmonicity), but also sequential cues (similarity to previous stimuli in the series) for auditory object formation. Although ORN was consistently elicited by two-object relative to one-object sounds, it was modulated by the presentation rate. With increasing probability of sound segregation, ORN amplitude decreased, and its scalp topography shifted from a bilateral towards a unilateral (left-hemispheric) activation pattern. Probability dependence of the ORN matches previous behavioral data. The differential sensitivity of the left- and right-hemispheric response to this manipulation may explain why an ORN probability effect was not obtained in previous electrophysiological studies. The specific functional role of ORN for concurrent sound perception was confirmed by dissociating it from the mismatch negativity (MMN) component as a correlate of processing sequential cues. ORN probability dependence is argued to be functionally relevant for the interaction of concurrent and sequential mechanisms of auditory scene analysis.

The dynamic attending theory as originally proposed by Jones, 1976. Psychol. Rev. 83(5), 323–355 posits that tone sequences presented at a regular rhythm entrain attentional oscillations and thereby facilitate the processing of sounds presented in phase with this rhythm. The increased interest in neural correlates of dynamic attending requires robust behavioral indicators of the phenomenon. Here we aimed to replicate and complement the most prominent experimental implementation of dynamic attending (Jones et al., 2002. Psychol. Sci. 13(4), 313–319). The paradigm uses a pitch comparison task in which two tones, the initial and the last of a longer series, have to be compared. In-between the two, distractor tones with variable pitch are presented, at a regular pace. A comparison tone presented in phase with the entrained rhythm is hypothesized to lead to better behavioral performance. Aiming for a conceptual replication, four different variations of the original paradigm were created which were followed by an exact replication attempt. Across all five experiments, only 40 of the 140 tested participants showed the hypothesized pattern of an inverted U-shaped profile in task accuracy, and the group average effects did not replicate the pattern reported by Jones et al., 2002. Psychol. Sci. 13(4), 313–319 in any of the five experiments. However, clear evidence for a relationship between musicality and overall behavioral performance was found. This study casts doubt on the suitability of the pitch comparison task for demonstrating auditory dynamic attending. We discuss alternative tasks that have been shown to support dynamic attending theory, thus lending themselves more readily to studying its neural correlates.This article is part of a Special Issue entitled SI: Prediction and Attention.

The ability to detect unexpected stimuli in the acoustic environment and determine their behavioral relevance to plan an appropriate reaction is critical for survival. This perspective article brings together several viewpoints and discusses current advances in understanding the mechanisms the auditory system implements to extract relevant information from incoming inputs and to identify unexpected events. This extraordinary sensitivity relies on the capacity to codify acoustic regularities, and is based on encoding properties that are present as early as the auditory midbrain. We review state-of-the-art studies on the processing of stimulus changes using non-invasive methods to record the summed electrical potentials in humans, and those that examine single-neuron responses in animal models. Human data will be based on mismatch negativity (MMN) and enhanced middle latency responses (MLR). Animal data will be based on the activity of single neurons at the cortical and subcortical levels, relating selective responses to novel stimuli to the MMN and to stimulus-specific neural adaptation (SSA). Theoretical models of the neural mechanisms that could create SSA and novelty responses will also be discussed.

While many studies have assessed the efficacy of similarity-based cues for auditory stream segregation, much less is known about whether and how the larger-scale structure of sound sequences support stream formation and the choice of sound organization. Two experiments investigated the effects of musical melody and rhythm on the segregation of two interleaved tone sequences. The two sets of tones fully overlapped in pitch range but differed from each other in interaural time and intensity. Unbeknownst to the listener, separately, each of the interleaved sequences was created from the notes of a different song. In different experimental conditions, the notes and/or their timing could either follow those of the songs or they could be scrambled or, in case of timing, set to be isochronous. Listeners were asked to continuously report whether they heard a single coherent sequence (integrated) or two concurrent streams (segregated). Although temporal overlap between tones from the two streams proved to be the strongest cue for stream segregation, significant effects of tonality and familiarity with the songs were also observed. These results suggest that the regular temporal patterns are utilized as cues in auditory stream segregation and that long-term memory is involved in this process.

Many sound sources emit signals in a predictable manner. The idea that predictability can be exploited to support the segregation of one source’s signal emissions from the overlapping signals of other sources has been expressed for a long time. Yet experimental evidence for a strong role of predictability within auditory scene analysis (ASA) has been scarce. Recently, there has been an upsurge in experimental and theoretical work on this topic resulting from fundamental changes in our perspective on how the brain extracts predictability from series of sensory events. Based on effortless predictive processing in the auditory system, it becomes more plausible that predictability would be available as a cue for sound source decomposition. In the present contribution, empirical evidence for such a role of predictability in ASA will be reviewed. It will be shown that predictability affects ASA both when it is present in the sound source of interest (perceptual foreground) and when it is present in other sound sources that the listener wishes to ignore (perceptual background). First evidence pointing towards age-related impairments in the latter capacity will be addressed. Moreover, it will be illustrated how effects of predictability can be shown by means of objective listening tests as well as by subjective report procedures, with the latter approach typically exploiting the multi-stable nature of auditory perception. Critical aspects of study design will be delineated to ensure that predictability effects can be unambiguously interpreted. Possible mechanisms for a functional role of predictability within ASA will be discussed, and an analogy with the old-plus-new heuristic for grouping simultaneous acoustic signals will be suggested.

In two experiments, we assessed the effects of combining different cues of concurrent sound segregation on the object-related negativity (ORN) and the P400 event-related potential components. Participants were presented with sequences of complex tones, half of which contained some manipulation: one or two harmonic partials were mistuned, delayed, or presented from a different location than the rest. In separate conditions, one, two, or three of these manipulations were combined. Participants watched a silent movie (passive listening) or reported after each tone whether they perceived one or two concurrent sounds (active listening). ORN was found in almost all conditions except for location difference alone during passive listening. Combining several cues or manipulating more than one partial consistently led to sub-additive effects on the ORN amplitude. These results support the view that ORN reflects a combined, feature-unspecific assessment of the auditory system regarding the contribution of two sources to the incoming sound.

Predictive accounts of perception have received increasing attention in the past 20 years. Detecting violations of auditory regularities, as reflected by the Mismatch Negativity (MMN) auditory event-related potential, is amongst the phenomena seamlessly fitting this approach. Largely based on the MMN literature, we propose a psychological conceptual framework called the Auditory Event Representation System (AERS), which is based on the assumption that auditory regularity violation detection and the formation of auditory perceptual objects are based on the same predictive regularity representations. Based on this notion, a computational model of auditory stream segregation, called CHAINS, has been developed. In CHAINS, the auditory sensory event representation of each incoming sound is considered for being the continuation of likely combinations of the preceding sounds in the sequence, thus providing alternative interpretations of the auditory input. Detecting repeating patterns allows predicting upcoming sound events, thus providing a test and potential support for the corresponding interpretation. Alternative interpretations continuously compete for perceptual dominance. In this paper, we briefly describe AERS and deduce some general constraints from this conceptual model. We then go on to illustrate how these constraints are computationally specified in CHAINS.

Many sound sources can only be recognised from the pattern of sounds they emit, and not from the individual sound events that make up their emission sequences. Auditory scene analysis addresses the difficult task of interpreting the sound world in terms of an unknown number of discrete sound sources (causes) with possibly overlapping signals, and therefore of associating each event with the appropriate source. There are potentially many different ways in which incoming events can be assigned to different causes, which means that the auditory system has to choose between them. This problem has been studied for many years using the auditory streaming paradigm, and recently it has become apparent that instead of making one fixed perceptual decision, given sufficient time, auditory perception switches back and forth between the alternatives—a phenomenon known as perceptual bi- or multi-stability. We propose a new model of auditory scene analysis at the core of which is a process that seeks to discover predictable patterns in the ongoing sound sequence. Representations of predictable fragments are created on the fly, and are maintained, strengthened or weakened on the basis of their predictive success, and conflict with other representations. Auditory perceptual organisation emerges spontaneously from the nature of the competition between these representations. We present detailed comparisons between the model simulations and data from an auditory streaming experiment, and show that the model accounts for many important findings, including: the emergence of, and switching between, alternative organisations; the influence of stimulus parameters on perceptual dominance, switching rate and perceptual phase durations; and the build-up of auditory streaming. The principal contribution of the model is to show that a two-stage process of pattern discovery and competition between incompatible patterns can account for both the contents (perceptual organisations) and the dynamics of human perception in auditory streaming., Author Summary The sound waves produced by objects in the environment mix together before reaching the ears. Before we can make sense of an auditory scene, our brains must solve the puzzle of how to disassemble the sound waveform into groupings that correspond to the original source signals. How is this feat accomplished? We propose that the auditory system continually scans the structure of incoming signals in search of clues to indicate which pieces belong together. For instance, sound events may belong together if they have similar features, or form part of a clear temporal pattern. However this process is complicated by lack of knowledge of future events and the many possible ways in which even a simple sound sequence can be decomposed. The biological solution is multistability: one possible interpretation of a sound is perceived initially, which then gives way to another interpretation, and so on. We propose a model of auditory multistability, in which fragmental descriptions of the signal compete and cooperate to explain the sound scene. We demonstrate, using simplified experimental stimuli, that the model can account for both the contents (perceptual organisations) and the dynamics of human perception in auditory streaming.

We tested whether incoming sounds are processed differently depending on how the preceding sound sequence has been interpreted by the brain. Sequences of a regularly repeating three-tone pattern, the perceived organization of which spontaneously switched back and forth between two alternative interpretations, were delivered to listeners. Occasionally, a regular tone was exchanged for a slightly or moderately lower one (deviants). The electroencephalogram (EEG) was recorded while listeners continuously marked their perception of the sound sequence. We found that for both the regular and the deviant tones, the early exogenous P1 and N1 amplitudes varied together with the perceived sound organization. Percept-dependent effects on the late endogenous N2 and P3a amplitudes were only found for deviant tones. These results suggest that the perceived sound organization affects sound processing both by modulating what information is extracted from incoming sounds as well as by influencing how deviant sound events are evaluated for further processing.

The regular behavior of sound sources helps us to make sense of the auditory environment. Regular patterns may, for instance, convey information on the identity of a sound source (such as the acoustic signature of a train moving on the rails). Yet typically, this signature overlaps in time with signals emitted from other sound sources. It is generally assumed that auditory regularity extraction cannot operate upon this mixture of signals because it only finds regularities between adjacent sounds. In this view, the auditory environment would be grouped into separate entities by means of readily available acoustic cues such as separation in frequency and location. Regularity extraction processes would then operate upon the resulting groups. Our new experimental evidence challenges this view. We presented two interleaved sound sequences which overlapped in frequency range and shared all acoustic parameters. The sequences only differed in their underlying regular patterns. We inserted deviants into one of the sequences to probe whether the regularity was extracted. In the first experiment, we found that these deviants elicited the mismatch negativity (MMN) component. Thus the auditory system was able to find the regularity between the non-adjacent sounds. Regularity extraction was not influenced by sequence cohesiveness as manipulated by the relative duration of tones and silent inter-tone-intervals. In the second experiment, we showed that a regularity connecting non-adjacent sounds was discovered only when the intervening sequence also contained a regular pattern, but not when the intervening sounds were randomly varying. This suggests that separate regular patterns are available to the auditory system as a cue for identifying signals coming from distinct sound sources. Thus auditory regularity extraction is not necessarily confined to a processing stage after initial sound grouping, but may precede grouping when other acoustic cues are unavailable.

The auditory system has been shown to detect predictability in a tone sequence, but does it use the extracted regularities for actually predicting the continuation of the sequence? The present study sought to find evidence for the generation of such predictions. Predictability was manipulated in an isochronous series of tones in which every other tone was a repetition of its predecessor. The existence of predictions was probed by occasionally omitting either the first (unpredictable) or the second (predictable) tone of a same-frequency tone pair. Event-related electrical brain activity elicited by the omission of an unpredictable tone differed from the response to the actual tone right from the tone onset. In contrast, early electrical brain activity elicited by the omission of a predictable tone was quite similar to the response to the actual tone. This suggests that the auditory system preactivates the neural circuits for expected input, using sequential predictions to specifically prepare for future acoustic events.

Traditional auditory oddball paradigms imply the brain's ability to encode regularities, but are not optimal for investigating the process of regularity establishment. In the present study, a dynamic experimental protocol was developed that simulates a more realistic auditory environment with changing regularities. The dynamic sequences were included in a distraction paradigm in order to study regularity extraction and application. Subjects discriminated the duration of sequentially presented tones. Without relevance to the task, tones repeated or changed in frequency according to a pattern unknown to the subject. When frequency repetitions were broken by a deviating tone, behavioral distraction (prolonged reaction time in the duration discrimination task) was elicited. Moreover, event-related brain potential components indicated deviance detection (mismatch negativity), involuntary attention switches (P3a), and attentional reorientation. These results suggest that regularities were extracted from the dynamic stimulation and were used to predict forthcoming stimuli. The effects were already observed with deviants occurring after as few as two presentations of a standard frequency, that is, violating a just emerging rule. Effects of regularity violation strengthened with the number of standard repetitions. Control stimuli comprising no regularity revealed that the observed effects were due to both improvements in standard processing (benefits of regularity establishment) and deteriorations in deviant processing (costs of regularity violation). Thus, regularities are exploited in two different ways: for an efficient processing of regularity-conforming events as well as for the detection of nonconforming, presumably important events. The present results underline the brain's flexibility in its adaptation to environmental demands.