Alexandra Bendixen

Early electrophysiological indicators for predictive processing in audition: a review.

The auditory system essentially deals with sequential type of input and thus requires processing that is particularly suited to extract stimulus relations within a sequence. Evidence from a variety of paradigms converges to show that the auditory system automatically uses stimulus predictability for facilitating its sequential processing. This type of predictive processing does not require attentional processing of the sounds or cognitive control of the predictions, nor does it involve the preparation of motor responses to the auditory stimuli. We will present a taxonomy of paradigms and resulting electrophysiological indicators for such automatic predictive processing in terms of event-related potential components and oscillatory activity. These indicators will include signals of fulfilled predictions (match signals such as N1 attenuation, repetition positivity, and early evoked gamma band response enhancement) as well as signals of violated predictions (mismatch signals such as the mismatch negativity and stimulus omission responses). We will show how recent approaches have revealed particularly early indicators of predictive processing down to the level of the auditory middle-latency responses. We will discuss the strength of the various indicators in terms of a truly predictive account of auditory processing (as opposed to, e.g., a retrospective verification of predictions). Finally, we will discuss the benefits of a predictive system within and beyond auditory processing. In conclusion, we argue in favor of the overwhelming evidence for predictions in audition, flexibly instantiated on different levels and timescales, and we aim to provide guidance along a variety of research paradigms illustrating the existence of these predictions.

Do N1/MMN, P3a, and RON form a strongly coupled chain reflecting the three stages of auditory distraction?

Distraction triggered by unexpected events is generally described in a serial model comprising (1) automatic detection of unexpected task-irrelevant events, (2) orienting towards the event, and (3) recovery from distraction. Processes taking place at the three stages are assumed to be reflected by the N1 and mismatch negativity (MMN); the P3a; and the reorienting negativity (RON) event-related potentials (ERPs), respectively. We investigated whether the processes indexed by these components form a strongly coupled chain, each co-varying with the preceding one. To this end, micro-sequence analysis of the ERPs elicited by unpredictable pitch-changes was conducted in an auditory duration discrimination task. Results indicated that the processes indexed by the above-mentioned ERPs are not strongly coupled. Pair-wise dissociations were found between the ERPs reflecting each processing stage: P3a can be elicited without concurrent N1-increase or MMN elicitation and without subsequent elicitation of the RON. Possible interpretations of P3a and RON are discussed.

I Heard That Coming: Event-Related Potential Evidence for Stimulus-Driven Prediction in the Auditory System

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.

Regularity Extraction and Application in Dynamic Auditory Stimulus Sequences

Traditional auditory oddball paradigms imply the brains 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 brains flexibility in its adaptation to environmental demands.

Multistability in auditory stream segregation: a predictive coding view

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.

Rapid extraction of auditory feature contingencies

Contingent relations between sensory events render the environment predictable and thus facilitate adaptive behavior. The human capacity to detect such relations has been comprehensively demonstrated in paradigms in which contingency rules were task-relevant or in which they applied to motor behavior. The extent to which contingencies can also be extracted from events that are unrelated to the current goals of the organism has remained largely unclear. The present study addressed the emergence of contingency-related effects for behaviorally irrelevant auditory stimuli and the cortical areas involved in the processing of such contingency rules. Contingent relations between different features of temporally separate events were embedded in a new dynamic protocol. Participants were presented with the auditory stimulus sequences while their attention was captured by a video. The mismatch negativity (MMN) component of the event-related brain potential (ERP) was employed as an electrophysiological correlate of contingency detection. MMN generators were localized by means of scalp current density (SCD) and primary current density (PCD) analyses with variable resolution electromagnetic tomography (VARETA). Results show that task-irrelevant contingencies can be extracted from about fifteen to twenty successive events conforming to the contingent relation. Topographic and tomographic analyses reveal the involvement of the auditory cortex in the processing of contingency violations. The present data provide evidence for the rapid encoding of complex extrapolative relations in sensory areas. This capacity is of fundamental importance for the organism in its attempt to model the sensory environment outside the focus of attention.

Processing of abstract rule violations in audition

The ability to encode rules and to detect rule-violating events outside the focus of attention is vital for adaptive behavior. Our brain recordings reveal that violations of abstract auditory rules are processed even when the sounds are unattended. When subjects performed a task related to the sounds but not to the rule, rule violations impaired task performance and activated a network involving supratemporal, parietal and frontal areas although none of the subjects acquired explicit knowledge of the rule or became aware of rule violations. When subjects tried to behaviorally detect rule violations, the brains automatic violation detection facilitated intentional detection. This shows the brains capacity for abstraction – an important cognitive function necessary to model the world. Our study provides the first evidence for the task-independence (i.e. automaticity) of this ability to encode abstract rules and for its immediate consequences for subsequent mental processes.

Neurophysiological evidence of impaired musical sound perception in cochlear-implant users

OBJECTIVE Music perception with a cochlear implant (CI) can be unsatisfactory because current-day implants are primarily designed to enable speech discrimination. The present study aimed at evaluating electrophysiological correlates of musical sound perception in CI users to help achieve the long-term goal of improved restoration of hearing in those individuals. METHODS Auditory discrimination accuracy in adult CI users (n=12) and matched normal-hearing controls (n=12) was measured by behavioral discrimination tasks and mismatch negativity (MMN) recordings. Discrimination profiles were obtained by using a set of clarinet sounds (original/vocoded) varying along different acoustic dimensions (frequency/intensity/duration) and deviation magnitudes (four levels). RESULTS Behavioral results and MMN recordings revealed reduced auditory discrimination accuracy in CI users. An inverse relationship was found between MMN amplitudes and duration of profound deafness. CONCLUSIONS CI users have difficulties in discriminating small changes in the acoustic properties of musical sounds. The recently developed multi-feature MMN paradigm (Pakarinen et al., 2007) can be used to objectively evaluate discrimination abilities of CI users for musical sounds. SIGNIFICANCE Measuring auditory discrimination functions by means of a multi-feature MMN paradigm could be of substantial clinical value by providing a comprehensive profile of the extent of restored hearing in CI users.

Hearing Silences: Human Auditory Processing Relies on Preactivation of Sound-Specific Brain Activity Patterns

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.

Memory trace formation for abstract auditory features and its consequences in different attentional contexts

The capacity for abstraction is vital for adaptive behavior. Based on behavioral data and event-related potentials (ERPs), the present study investigates the brains ability to encode abstract auditory rules with a dynamic approach in which rules constantly emerge and vanish. In successive conditions, abstract rules are task-irrelevant and task-relevant. Results show that as few as two consecutive exemplars of an abstract feature (frequency relation between successive tones) are sufficient for rule extraction. The extraction of just emerging rules is independent of the amount of attention devoted to the rules, and it is not modulated by further rule-conforming exemplars. The extracted rules are immediately applied, as evidenced by the interference of task-irrelevant rule violations with concurrent mental processes (distraction condition) and by the conscious detection of task-relevant violations (detection condition). The ability to rapidly encode abstract rules and to detect presumably important rule-violating events underlines the brains adaptability to the environmental demands.