Suppression and omission effects in auditory predictive processing-Two of the same?

Recent theories describe perception as an inferential process based on internal predictive models that are adjusted by prediction violations (prediction error). Two different modulations of the auditory N1 event-related brain potential component are often discussed as an expression of auditory predictive processing. The sound-related N1 component is attenuated for self-generated sounds compared to the N1 elicited by externally generated sounds (N1 suppression). An omission-related component in the N1 time-range is elicited when the self-generated sounds are occasionally omitted (omission N1). Both phenomena were explained by action-related forward modelling, which takes place when the sensory input is predictable: prediction error signals are reduced when predicted sensory input is presented (N1 suppression) and elicited when predicted sensory input is omitted (omission N1). This common theoretical account is appealing but has not yet been directly tested. We manipulated the predictability of a sound in a self-generation paradigm in which, in two conditions, either 80% or 50% of the button presses did generate a sound, inducing a strong or a weak expectation for the occurrence of the sound. Consistent with the forward modelling account, an omission N1 was observed in the 80% but not in the 50% condition. However, N1 suppression was highly similar in both conditions. Thus, our results demonstrate a clear effect of predictability for the omission N1 but not for the N1 suppression. These results imply that the two phenomena rely (at least in part) on different mechanisms and challenge prediction related accounts of N1 suppression.

Conditional deviant repetition in the oddball paradigm modulates processing at the level of P3a but not MMN.

The auditory system has an amazing ability to rapidly encode auditory regularities. Evidence comes from the popular oddball paradigm, in which frequent (standard) sounds are occasionally exchanged for rare deviant sounds, which then elicit signs of prediction error based on their unexpectedness (e.g., MMN and P3a). Here, we examine the widely neglected characteristics of deviants being bearers of predictive information themselves; naive participants listened to sound sequences constructed according to a new, modified version of the oddball paradigm including two types of deviants that followed diametrically opposed rules: one deviant sound occurred mostly in pairs (repetition rule), the other deviant sound occurred mostly in isolation (non-repetition rule). Due to this manipulation, the sound following a first deviant (either the same deviant or a standard) was either predictable or unpredictable based on its conditional probability associated with the preceding deviant sound. Our behavioral results from an active deviant detection task replicate previous findings that deviant repetition rules (based on conditional probability) can be extracted when behaviorally relevant. Our electrophysiological findings obtained in a passive listening setting indicate that conditional probability also translates into differential processing at the P3a level. However, MMN was confined to global deviants and was not sensitive to conditional probability. This suggests that higher-level processing concerned with stimulus selection and/or evaluation (reflected in P3a) but not lower-level sensory processing (reflected in MMN) considers rarely encountered rules.

Mechanisms of sustained perceptual entrainment after stimulus offset.

Temporal alignment of neural activity to rhythmic stimulation has been suggested to result from a resonating internal neural oscillator mechanism, but can also be explained by interval-based temporal prediction. Here, we investigate behavioural and brain responses in the post-stimulation period to compare an oscillatory versus an interval-based account. Hickok et al.'s (2015) behavioural paradigm yielded results that relate to a neural oscillatory entrainment mechanism. We adapted the paradigm to an event-related potential (ERP) suitable design: a periodic sequence was followed, in half of the trials, by near-threshold targets embedded in noise. The targets were played in various phases in relation to the preceding sequences' period. Participants had to detect whether targets were played or not, and their EEG was recorded. Both behavioural results and the P300 component of the ERP were not only partially consistent with an oscillatory mechanism but also partially consistent with an interval-based attentional gain mechanism. Instead, data obtained in the post-entrainment period can best be explained with a combination of both mechanisms.

Neural signatures of automatic repetition detection in temporally regular and jittered acoustic sequences.

Detection of repeating patterns within continuous sound streams is crucial for efficient auditory perception. Previous studies demonstrated a remarkable sensitivity of the human auditory system to periodic repetitions in unfamiliar, meaningless sounds. Automatic repetition detection was reflected in different EEG markers, including sustained activity, neural synchronisation, and event-related responses to pattern occurrences. The current study investigated how listeners' attention and the temporal regularity of a sound modulate repetition perception, and how this influence is reflected in different EEG markers that were previously suggested to subserve dissociable functions. We reanalysed data of a previous study in which listeners were presented with sequences of unfamiliar artificial sounds that either contained repetitions of a certain sound segment or not. Repeating patterns occurred either regularly or with a temporal jitter within the sequences, and participants' attention was directed either towards the pattern repetitions or away from the auditory stimulation. Across both regular and jittered sequences during both attention and in-attention, pattern repetitions led to increased sustained activity throughout the sequence, evoked a characteristic positivity-negativity complex in the event-related potential, and enhanced inter-trial phase coherence of low-frequency oscillatory activity time-locked to repeating pattern onsets. While regularity only had a minor (if any) influence, attention significantly strengthened pattern repetition perception, which was consistently reflected in all three EEG markers. These findings suggest that the detection of pattern repetitions within continuous sounds relies on a flexible mechanism that is robust against in-attention and temporal irregularity, both of which typically occur in naturalistic listening situations. Yet, attention to the auditory input can enhance processing of repeating patterns and improve repetition detection.

Markov chains as a proxy for the predictive memory representations underlying mismatch negativity.

Events not conforming to a regularity inherent to a sequence of events elicit prediction error signals of the brain such as the Mismatch Negativity (MMN) and impair behavioral task performance. Events conforming to a regularity lead to attenuation of brain activity such as stimulus-specific adaptation (SSA) and behavioral benefits. Such findings are usually explained by theories stating that the information processing system predicts the forthcoming event of the sequence via detected sequential regularities. A mathematical model that is widely used to describe, to analyze and to generate event sequences are Markov chains: They contain a set of possible events and a set of probabilities for transitions between these events (transition matrix) that allow to predict the next event on the basis of the current event and the transition probabilities. The accuracy of such a prediction depends on the distribution of the transition probabilities. We argue that Markov chains also have useful applications when studying cognitive brain functions. The transition matrix can be regarded as a proxy for generative memory representations that the brain uses to predict the next event. We assume that detected regularities in a sequence of events correspond to (a subset of) the entries in the transition matrix. We apply this idea to the Mismatch Negativity (MMN) research and examine three types of MMN paradigms: classical oddball paradigms emphasizing sound probabilities, between-sound regularity paradigms manipulating transition probabilities between adjacent sounds, and action-sound coupling paradigms in which sounds are associated with actions and their intended effects. We show that the Markovian view on MMN yields theoretically relevant insights into the brain processes underlying MMN and stimulates experimental designs to study the brain's processing of event sequences.

Deviants violating higher-order auditory regularities can become predictive and facilitate behaviour.

The human auditory system is believed to represent regularities inherent in auditory information in internal models. Sounds not matching the standard regularity (deviants) elicit prediction error, alerting the system to information not explainable within currently active models. Here, we examine the widely neglected characteristic of deviants bearing predictive information themselves. In a modified version of the oddball paradigm, using higher-order regularities, we set up different expectations regarding the sound following a deviant. Higher-order regularities were defined by the relation of pitch within tone pairs (rather than absolute pitch of individual tones). In a deviant detection task participants listened to oddball sequences including two deviant types following diametrically opposed rules: one occurred mostly in succession (high repetition probability) and the other mostly in isolation (low repetition probability). Participants in Experiment 1 were not informed (naïve), whereas in Experiment 2 they were made aware of the repetition rules. Response times significantly decreased from first to second deviant when repetition probability was high-albeit more in the presence of explicit rule knowledge. There was no evidence of a facilitation effect when repetition probability was low. Significantly more false alarms occurred in response to standards following high compared with low repetition probability deviants, but only in participants aware of the repetition rules. These findings provide evidence that not only deviants violating lower- but also higher-order regularities can inform predictions about auditory events. More generally, they confirm the utility of this new paradigm to gather further insights into the predictive properties of the human brain.

Perceptual learning of random acoustic patterns: Impact of temporal regularity and attention.

Perceptual learning is a powerful mechanism to enhance perceptual abilities and to form robust memory representations of previously unfamiliar sounds. Memory formation through repeated exposure takes place even for random and complex acoustic patterns devoid of semantic content. The current study sought to scrutinise how perceptual learning of random acoustic patterns is shaped by two potential modulators: temporal regularity of pattern repetition and listeners' attention. To this end, we adapted an established implicit learning paradigm and presented short acoustic sequences that could contain embedded repetitions of a certain sound segment (i.e., pattern) or not. During each experimental block, one repeating pattern recurred across multiple trials, whereas the other patterns were presented in only one trial. During the presentation of sound sequences that contained either temporally regular or jittered within-trial pattern repetitions, participants' attention was directed either towards or away from the auditory stimulation. Overall, we found a memory-related modulation of the event-related potential (ERP) and an increase in inter-trial phase coherence for patterns that recurred across multiple trials (compared to non-recurring patterns), accompanied by a performance increase in a (within-trial) repetition detection task when listeners attended the sounds. Remarkably, we show a memory-related ERP effect even for the first pattern occurrence per sequence when participants attended the sounds, but not when they were engaged in a visual distractor task. These findings suggest that learning of unfamiliar sound patterns is robust against temporal irregularity and inattention, but attention facilitates access to established memory representations upon first occurrence within a sequence.

Auditory representations for long lasting sounds: Insights from event-related brain potentials and neural oscillations.

The basic features of short sounds, such as frequency and intensity including their temporal dynamics, are integrated in a unitary representation. Knowledge on how our brain processes long lasting sounds is scarce. We review research utilizing the Mismatch Negativity event-related potential and neural oscillatory activity for studying representations for long lasting simple versus complex sounds such as sinusoidal tones versus speech. There is evidence for a temporal constraint in the formation of auditory representations: Auditory edges like sound onsets within long lasting sounds open a temporal window of about 350 ms in which the sounds' dynamics are integrated into a representation, while information beyond that window contributes less to that representation. This integration window segments the auditory input into short chunks. We argue that the representations established in adjacent integration windows can be concatenated into an auditory representation of a long sound, thus, overcoming the temporal constraint.

Within- and between-subject consistency of perceptual segmentation in periodic noise: A combined behavioral tapping and EEG study.

It is remarkable that human listeners can perceive periodicity in noise, as the isochronous repetition of a particular noise segment is not accompanied by salient physical cues in the acoustic signal. Previous research suggested that listeners rely on short temporally local and idiosyncratic features to perceptually segment periodic noise sequences. The present study sought to test this assumption by disentangling consistency of perceptual segmentation within and between listeners. Presented periodic noise sequences either consisted of seamless repetitions of a 500-ms segment or of repetitions of a 200-ms segment that were interleaved with 300-ms portions of random noise. Both within- and between-subject consistency was stronger for interleaved (compared with seamless) periodic sequences. The increased consistency likely resulted from reduced temporal jitter of potential features used for perceptual segmentation when the recurring segment was shorter and occurred interleaved with random noise. These results support the notion that perceptual segmentation of periodic noise relies on subtle temporally local features. However, the finding that some specific noise sequences were segmented more consistently across listeners than others challenges the assumption that the features are necessarily idiosyncratic. Instead, in some specific noise samples, a preference for certain spectral features is shared between individuals.

A link between age, affect, and predictions?

The prevalence of depressive symptoms decreases from late adolescence to middle age adulthood. Furthermore, despite significant losses in motor and cognitive functioning, overall emotional well-being tends to increase with age, and a bias to positive information has been observed multiple times. Several causes have been discussed for this age-related development, such as improvement in emotion regulation, less regret, and higher socioeconomic status. Here, we explore a further explanation. Our minds host mental models that generate predictions about forthcoming events to successfully interact with our physical and social environment. To keep these models faithful, the difference between the predicted and the actual event, that is, the prediction error, is computed. We argue that prediction errors are attenuated in the middle age and older mind, which, in turn, may translate to less negative affect, lower susceptibility to affective disorders, and possibly, to a bias to positive information. Our proposal is primarily linked to perceptual inferences, but may hold as well for higher-level, cognitive, and emotional forms of error processing.

Intention-based predictive information modulates auditory deviance processing.

The human brain is highly responsive to (deviant) sounds violating an auditory regularity. Respective brain responses are usually investigated in situations when the sounds were produced by the experimenter. Acknowledging that humans also actively produce sounds, the present event-related potential study tested for differences in the brain responses to deviants that were produced by the listeners by pressing one of two buttons. In one condition, deviants were unpredictable with respect to the button-sound association. In another condition, deviants were predictable with high validity yielding correctly predicted deviants and incorrectly predicted (mispredicted) deviants. Temporal principal component analysis revealed deviant-specific N1 enhancement, mismatch negativity (MMN) and P3a. N1 enhancements were highly similar for each deviant type, indicating that the underlying neural mechanism is not affected by intention-based expectation about the self-produced forthcoming sound. The MMN was abolished for predictable deviants, suggesting that the intention-based prediction for a deviant can overwrite the prediction derived from the auditory regularity (predicting a standard). The P3a was present for each deviant type but was largest for mispredicted deviants. It is argued that the processes underlying P3a not only evaluate the deviant with respect to the fact that it violates an auditory regularity but also with respect to the intended sensorial effect of an action. Overall, our results specify current theories of auditory predictive processing, as they reveal that intention-based predictions exert different effects on different deviance-specific brain responses.

Action effect predictions in 'what', 'when', and 'whether' intentional actions.

It has been proposed that intentional action can be separated into three major types depending on the nature of the action choice - what (selecting what to do), when (selecting when to act) and whether (to perform the action or not). While many theories on action control assume that intentional action involves the prediction of action effects, there has not been any attempt to compare the three types of intentional actions (what, when, whether) with respect to action-effect prediction. Here, we employ an action-effect prediction paradigm where participants select the action on every trial based on either the what (choosing between alternative actions), when (choosing to respond at different time points) or whether (choosing to perform an action or not) action components, and each action choice is followed by either a predicted (standard) or a mispredicted (deviant) tone. We found a significant P2 difference between standard/deviant tones reflecting the formation of action-effect predictions regardless of whether the action choice was based on the 'what', 'when' or 'whether' decision. Furthermore, our analysis revealed that this P2 difference for the prediction effect was not observable in non-action trials within the 'whether' condition, which suggests an action-specific prediction process.

Hearing "Birch" Hampers Saying "Duck"-An Event-Related Potential Study on Phonological Interference in Immediate and Delayed Word Production.

When speakers name a picture (e.g., "duck"), a distractor word phonologically related to an alternative name (e.g., "birch" related to "bird") slows down naming responses compared with an unrelated distractor word. This interference effect obtained with the picture-word interference task is assumed to reflect the phonological coactivation of close semantic competitors and is critical for evaluating contemporary models of word production. In this study, we determined the ERP signature of this effect in immediate and delayed versions of the picture-word interference task. ERPs revealed a differential processing of related and unrelated distractors: an early (305-436 msec) and a late (537-713 msec) negativity for related as compared with unrelated distractors. In the behavioral data, the interference effect was only found in immediate naming, whereas its ERP signature was also present in delayed naming. The time window of the earlier ERP effect suggests that the behavioral interference effect indeed emerges at a phonological processing level, whereas the functional significance of the later ERP effect is as yet not clear. The finding of a robust ERP correlate of phonological coactivation might facilitate future research on lexical processing in word production.

Neural entrainment via perceptual inferences.

Entrainment depends on sequential neural phase reset by regular stimulus onset, a temporal parameter. Entraining to sequences of identical stimuli also entails stimulus feature predictability, but this component is not readily separable from temporal regularity. To test if spectral regularities concur with temporal regularities in determining the strength of auditory entrainment, we devised sound sequences that varied in conditional perceptual inferences based on deviant sound repetition probability: strong inference (100% repetition probability: If a deviant appears, then it will repeat), weak inference (75% repetition probability) and no inference (50%: A deviant may or may not repeat with equal probability). We recorded EEG data from 15 young human participants pre-attentively listening to the experimental sound sequences delivered either isochronously or anisochronously (±20% jitter), at both delta (1.67 Hz) and theta (6.67 Hz) stimulation rates. Strong perceptual inferences significantly enhanced entrainment at either stimulation rate and determined positive correlations between precision in phase distribution at the onset of deviant trials and entrained power. We conclude that both spectral predictability and temporal regularity govern entrainment via neural phase control.

The auditory brain in action: Intention determines predictive processing in the auditory system-A review of current paradigms and findings.

According to the ideomotor theory, action may serve to produce desired sensory outcomes. Perception has been widely described in terms of sensory predictions arising due to top-down input from higher order cortical areas. Here, we demonstrate that the action intention results in reliable top-down predictions that modulate the auditory brain responses. We bring together several lines of research, including sensory attenuation, active oddball, and action-related omission studies: Together, the results suggest that the intention-based predictions modulate several steps in the sound processing hierarchy, from preattentive to evaluation-related processes, also when controlling for additional prediction sources (i.e., sound regularity). We propose an integrative theoretical framework-the extended auditory event representation system (AERS), a model compatible with the ideomotor theory, theory of event coding, and predictive coding. Initially introduced to describe regularity-based auditory predictions, we argue that the extended AERS explains the effects of action intention on auditory processing while additionally allowing studying the differences and commonalities between intention- and regularity-based predictions-we thus believe that this framework could guide future research on action and perception.

The posterior auditory field is the chief generator of prediction error signals in the auditory cortex.

The auditory cortex (AC) encompasses distinct fields subserving partly different aspects of sound processing. One essential function of the AC is the detection of unpredicted sounds, as revealed by differential neural activity to predictable and unpredictable sounds. According to the predictive coding framework, this effect can be explained by repetition suppression and/or prediction error signaling. The present study investigates functional specialization of the rat AC fields in repetition suppression and prediction error by combining a tone frequency oddball paradigm (involving high-probable standard and low-probable deviant tones) with two different control sequences (many-standards and cascade). Tones in the control sequences were comparable to deviant events with respect to neural adaptation but were not violating a regularity. Therefore, a difference in the neural activity between deviant and control tones indicates a prediction error effect, whereas a difference between control and standard tones indicates a repetition suppression effect. Single-unit recordings revealed by far the largest prediction error effects for the posterior auditory field, while the primary auditory cortex, the anterior auditory field, the ventral auditory field, and the suprarhinal auditory field were dominated by repetition suppression effects. Statistically significant repetition suppression effects occurred in all AC fields, whereas prediction error effects were less robust in the primary auditory cortex and the anterior auditory field. Results indicate that the non-lemniscal, posterior auditory field is more engaged in context-dependent processing underlying deviance-detection than the other AC fields, which are more sensitive to stimulus-dependent effects underlying differential degrees of neural adaptation.

Auditory Pattern Representations Under Conditions of Uncertainty-An ERP Study.

The auditory system is able to recognize auditory objects and is thought to form predictive models of them even though the acoustic information arriving at our ears is often imperfect, intermixed, or distorted. We investigated implicit regularity extraction for acoustically intact versus disrupted six-tone sound patterns via event-related potentials (ERPs). In an exact-repetition condition, identical patterns were repeated; in two distorted-repetition conditions, one randomly chosen segment in each sound pattern was replaced either by white noise or by a wrong pitch. In a roving-standard paradigm, sound patterns were repeated 1-12 times (standards) in a row before a new pattern (deviant) occurred. The participants were not informed about the roving rule and had to detect rarely occurring loudness changes. Behavioral detectability of pattern changes was assessed in a subsequent behavioral task. Pattern changes (standard vs. deviant) elicited mismatch negativity (MMN) and P3a, and were behaviorally detected above the chance level in all conditions, suggesting that the auditory system extracts regularities despite distortions in the acoustic input. However, MMN and P3a amplitude were decreased by distortions. At the level of MMN, both types of distortions caused similar impairments, suggesting that auditory regularity extraction is largely determined by the stimulus statistics of matching information. At the level of P3a, wrong-pitch distortions caused larger decreases than white-noise distortions. Wrong-pitch distortions likely prevented the engagement of restoration mechanisms and the segregation of disrupted from true pattern segments, causing stronger informational interference with the relevant pattern information.

Attentional Processing of Disgust and Fear and Its Relationship With Contamination-Based Obsessive-Compulsive Symptoms: Stronger Response Urgency to Disgusting Stimuli in Disgust-Prone Individuals.

Disgust has recently been characterized as a low-urgency emotion, particularly compared to fear. The aim of the present study is to clarify whether behavioral inhibition during disgust engagement is characteristic of a low-urgency emotion and thus indicates self-imposed attentional avoidance in comparison to fear. Therefore, 54 healthy participants performed an emotional go/no-go task with disgust- and fear-relevant as well as neutral pictures. Furthermore, heart rate activity and facial muscle activity on the fear-specific m. corrugator supercilli and the disgust-specific m. levator labii were assessed. The results partially support the temporal urgency hypothesis of disgust. The emotion conditions significantly differed in emotional engagement and in the facial muscle activity of the m. levator labii as expected. However, contrary to our expectations, no differences between the emotion conditions regarding behavioral inhibition as well as heart rate change could be found. Furthermore, individuals with a higher-trait disgust proneness showed faster reactions and higher activity of the m. levator labii in response to disgust stimuli. The results show that different trait levels influence attentional engagement and physiological parameters but have only a small effect on behavioral inhibition.

Word class and word frequency in the MMN looking glass.

The effects of lexical meaning and lexical familiarity on auditory deviance detection were investigated by presenting oddball sequences of words, while participants ignored the stimuli. Stimulus sequences were composed of words that were varied in word class (nouns vs. functions words) and frequency of language use (high vs. low frequency) in a factorial design with the roles of frequently presented stimuli (Standards) and infrequently presented ones (Deviants) were fully crossed. Deviants elicited the Mismatch Negativity component of the event-related brain potential. Modulating effects of lexical meaning were obtained, revealing processing advantages for denotationally meaningful items. However, no effect of word frequency was observed. These results demonstrate that an apparently low-level function, such as auditory deviance detection utilizes information from the mental lexicon even for task-irrelevant stimuli.