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.

Somatosensory omissions reveal action-related predictive processing.

The intricate relation between action and somatosensory perception has been studied extensively in the past decades. Generally, a forward model is thought to predict the somatosensory consequences of an action. These models propose that when an action is reliably coupled to a tactile stimulus, unexpected absence of the stimulus should elicit prediction error. Although such omission responses have been demonstrated in the auditory modality, it remains unknown whether this mechanism generalizes across modalities. This study therefore aimed to record action-induced somatosensory omission responses using EEG in humans. Self-paced button presses were coupled to somatosensory stimuli in 88% of trials, allowing a prediction, or in 50% of trials, not allowing a prediction. In the 88% condition, stimulus omission resulted in a neural response consisting of multiple components, as revealed by temporal principal component analysis. The oN1 response suggests similar sensory sources as stimulus-evoked activity, but an origin outside primary cortex. Subsequent oN2 and oP3 responses, as previously observed in the auditory domain, likely reflect modality-unspecific higher order processes. Together, findings straightforwardly demonstrate somatosensory predictions during action and provide evidence for a partially amodal mechanism of prediction error generation.

On the relationship of arousal and attentional distraction by emotional novel sounds.

Unexpected and task-irrelevant sounds can impair performance in a task. It has been shown that highly arousing emotional distractor sounds impaired performance less compared to moderately arousing neutral distractor sounds. The present study tests whether these differential emotion-related distraction effects are directly related to an enhancement of arousal evoked by processing of emotional distractor sounds. We disentangled costs of orienting of attention and benefits of increased arousal levels during the presentation of highly arousing emotional and moderately arousing neutral novel sounds that were embedded in a sequence of repeated standard sounds. We used sound-related pupil dilation responses as a marker of arousal and RTs as a marker of distraction in a visual categorization task in 57 healthy young adults. Multilevel analyses revealed increased RT and increased pupil dilation in response to novel vs. standard sounds. Emotional novel sounds reduced distraction effects on the behavioral level and increased pupil dilation responses compared to neutral novel sounds. Bayes Factors revealed strong evidence against an inverse proportional relationship between behavioral distraction effects and sound-related pupil dilation responses for emotional sounds. Given that the activity of the locus coeruleus has been linked to both changes in pupil diameter and arousal, it may embody an indirect relationship as a common antecedent by the release of norepinephrine into brain networks involved in attention control and control of the pupil. The present study provides new insights into the relation of changes in arousal and attentional distraction during the processing of emotional task-irrelevant novel sounds.

Salient omissions-pupil dilation in response to unexpected omissions of sound and touch.

Introduction: Recent theories describe perception as an inferential process based on internal predictive models adjusted by means of prediction violations (prediction error). To study and demonstrate predictive processing in the brain the use of unexpected stimulus omissions has been suggested as a promising approach as the evoked brain responses are uncontaminated by responses to stimuli. Here, we aimed to investigate the pupil's response to unexpected stimulus omissions in order to better understand surprise and orienting of attention resulting from prediction violation. So far only few studies have used omission in pupillometry research and results have been inconsistent. Methods: This study adapted an EEG paradigm that has been shown to elicit omission responses in auditory and somatosensory modalities. Healthy adults pressed a button at their own pace, which resulted in the presentation of sounds or tactile stimuli in either 88%, 50% or 0% (motor-control) of cases. Pupil size was recorded continuously and averaged to analyze the pupil dilation response associated with each condition. Results: Results revealed that omission responses were observed in both modalities in the 88%-condition compared to motor-control. Similar pupil omission responses were observed between modalities, suggesting modality-unspecific activation of the underlying brain circuits. Discussion: In combination with previous omission studies using EEG, the findings demonstrate predictive models in brain processing and point to the involvement of subcortical structures in the omission response. Our pupillometry approach is especially suitable to study sensory prediction in vulnerable populations within the psychiatric field.

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.

Attentional control in middle childhood is highly dynamic-Strong initial distraction is followed by advanced attention control.

The ability to shield against distraction while focusing on a task requires the operation of executive functions and is essential for successful learning. We investigated the short-term dynamics of distraction control in a data set of 269 children aged 4-10 years and 51 adults pooled from three studies using multilevel models. Participants performed a visual categorization task while a task-irrelevant sequence of sounds was presented which consisted of frequently repeated standard sounds and rarely interspersed novel sounds. On average, participants responded slower in the categorization task after novel sounds. This distraction effect was more pronounced in children. Throughout the experiment, the initially strong distraction effects declined to the level of adults in the groups of 6- to 10-year-olds. Such a decline was neither observed in the groups of the 4- and 5-year-olds, who consistently showed a high level of distraction, nor in adults, who showed a constantly low level of distraction throughout the experimental session. Results indicate that distraction control is a highly dynamic process that qualitatively and quantitatively differs between age groups. We conclude that the analysis of short-term dynamics provides valuable insights into the development of attention control and might explain inconsistent findings regarding distraction control in middle childhood. In addition, models of attention control need to be refined to account for age-dependent rapid learning mechanisms. Our findings have implications for the design of learning situations and provide an additional source of information for the diagnosis and treatment of children with attention deficit disorders.

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.

The effect of background speech on attentive sound processing: A pupil dilation study.

Listening to task-irrelevant speech while performing a cognitive task can involuntarily deviate our attention and lead to decreases in performance. One explanation for the impairing effect of irrelevant speech is that semantic processing can consume attentional resources. In the present study, we tested this assumption by measuring performance in a non-linguistic attentional task while participants were exposed to meaningful (native) and non-meaningful (foreign) speech. Moreover, based on the tight relation between pupillometry and attentional processes, we also registered changes in pupil diameter size to quantify the effect of meaningfulness upon attentional allocation. To these aims, we recruited 41 native German speakers who had neither received formal instruction in French nor had extensive informal contact with this language. The focal task consisted of an auditory oddball task. Participants performed a duration discrimination task containing frequently repeated standard sounds and rarely presented deviant sounds while a story was read in German or (non-meaningful) French in the background. Our results revealed that, whereas effects of language meaningfulness on attention were not detectable at the behavioural level, participants' pupil dilated more in response to the sounds of the auditory task when background speech was played in non-meaningful French compared to German, independent of sound type. In line with the initial hypothesis, this suggested that semantic processing of the native language required attentional resources, which lead to fewer resources devoted to the processing of the sounds of the focal task. Our results highlight the potential of the pupil dilation response for the investigation of subtle cognitive processes that might not surface when only behaviour is measured.

A tutorial on the use of temporal principal component analysis in developmental ERP research - Opportunities and challenges.

Developmental researchers are often interested in event-related potentials (ERPs). Data-analytic approaches based on the observed ERP suffer from major problems such as arbitrary definition of analysis time windows and regions of interest and the observed ERP being a mixture of latent underlying components. Temporal principal component analysis (PCA) can reduce these problems. However, its application in developmental research comes with the unique challenge that the component structure differs between age groups (so-called measurement non-invariance). Separate PCAs for the groups can cope with this challenge. We demonstrate how to make results from separate PCAs accessible for inferential statistics by re-scaling to original units. This tutorial enables readers with a focus on developmental research to conduct a PCA-based ERP analysis of amplitude differences. We explain the benefits of a PCA-based approach, introduce the PCA model and demonstrate its application to a developmental research question using real-data from a child and an adult group (code and data openly available). Finally, we discuss how to cope with typical challenges during the analysis and name potential limitations such as suboptimal decomposition results, data-driven analysis decisions and latency shifts.

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.

Sound omission related brain responses in children.

Action is an important way for children to learn about the world. Recent theories suggest that action is inherently accompanied by the sensory prediction of its effects. Such predictions can be revealed by rarely omitting the expected sensory consequence of the action, resulting in an omission response that is observable in the EEG. Although prediction errors play an important role in models of learning and development, little is known about omission-related brain responses in children. This study used a motor-auditory omission paradigm, testing a group of 6-8-year-old children and an adult group (N = 31 each). In an identity-specific condition, the sound coupled to the motor action was predictable, while in an identity unspecific condition the sound was unpredictable. Results of a temporal principal component analysis revealed that sound-related brain responses underlying the N1-complex differed considerably between age groups. Despite these developmental differences, omission responses (oN1) were similar between age groups. Two subcomponents of the oN1 were differently affected by specific and unspecific predictions. Results demonstrate that children, independent from the maturation of sound processing mechanisms, can implement specific and unspecific predictions as flexibly as adults. This supports theories that regard action and prediction error as important drivers of cognitive development.

Tablet PC use directly affects children's perception and attention.

Children currently grow up with a marked increase in interactive digital mobile media. To what extent digital media directly modulate children's perception and attention is largely unknown. We investigated the processing of task-irrelevant auditory information while 37 children aged 6;8-9;1-years played the identical card game on a tablet PC or with the experimenter in reality. The sound sequence included repeated standard sounds and occasionally novel sounds. Event-related potentials in the EEG, that reflect sound-related processes of perception and attention, were measured. Sounds evoked increased amplitudes of the ERP components P1, P2 and P3a during the interaction with the tablet PC compared to the human interaction. This indicates enhanced early processing of task-irrelevant information and increased allocation of attention to sounds throughout the interaction with a tablet PC compared to a human partner. Results suggest direct effects of typical situations, where children interact with a tablet PC, on neuronal mechanisms that drive perception and attention in the developing brain. More research into this phenomena is required to make specific suggestions for developing digital interactive learning programs.

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.

The encoding of stochastic regularities is facilitated by action-effect predictions.

Our brains continuously build and update predictive models of the world, sources of prediction being drawn for example from sensory regularities and/or our own actions. Yet, recent results in the auditory system indicate that stochastic regularities may not be easily encoded when a rare medium pitch deviant is presented between frequent high and low pitch standard sounds in random order, as reflected in the lack of sensory prediction error event-related potentials [i.e., mismatch negativity (MMN)]. We wanted to test the implication of the predictive coding theory that predictions based on higher-order generative models-here, based on action intention, are fed top-down in the hierarchy to sensory levels. Participants produced random sequences of high and low pitch sounds by button presses in two conditions: In a "specific" condition, one button produced high and the other low pitch sounds; in an "unspecific" condition, both buttons randomly produced high or low-pitch sounds. Rare medium pitch deviants elicited larger MMN and N2 responses in the "specific" compared to the "unspecific" condition, despite equal sound probabilities. These results thus demonstrate that action-effect predictions can boost stochastic regularity-based predictions and engage higher-order deviance detection processes, extending previous notions on the role of action predictions at sensory levels.

Cross-modal predictive processing depends on context rather than local contingencies.

Visual symbols or events may provide predictive information on to-be-expected sound events. When the perceived sound does not confirm the visual prediction, the incongruency response (IR), a prediction error signal of the event-related brain potentials, is elicited. It is unclear whether predictions are derived from lower-level local contingencies (e.g., recent events or repetitions) or from higher-level global rules applied top-down. In a recent study, sound pitch was predicted by a preceding note symbol. IR elicitation was confined to the condition where one of two sounds was presented more frequently and was not present with equal probability of both sounds. These findings suggest that local repetitions support predictive cross-modal processing. On the other hand, IR has also been observed with equal stimulus probabilities, where visual patterns predicted the upcoming sound sequence. This suggests the application of global rules. Here, we investigated the influence of stimulus repetition on the elicitation of the IR by presenting identical trial trains of a particular visual note symbol cueing a particular sound resulting either in a congruent or an incongruent pair. Trains of four different lengths: 1, 2, 4, or 7 were presented. The IR was observed already after a single presentation of a congruent visual-cue-sound combination and did not change in amplitude as trial train length increased. We conclude that higher-level associations applied in a top-down manner are involved in elicitation of the prediction error signal reflected by the IR, independent from local contingencies.

Distraction of attention by novel sounds in children declines fast.

New task-irrelevant sounds can distract attention. This study specifies the impact of stimulus novelty and of learning on attention control in three groups of children aged 6-7, 8, and 9-10 years and an adult control group. Participants (N = 179) were instructed to ignore a sound sequence including standard sounds and novel or repeated distractor sounds, while performing a visual categorization task. Distractor sounds impaired performance in children more than in adult controls, demonstrating the long-term development of attention control. Children, but not adults, were more distracted by novel than by repeated sounds, indicating increased sensitivity to novel information. Children, in particular younger children, were highly distracted during the first presentations of novel sounds compared to adults, while no age differences were observed for the last presentations. Results highlight the age-related impact of auditory novel information on attention and characterize the rapid development of attention control mechanisms as a function of age and exposure to irrelevant novel sounds.

Modulation of early auditory processing by visual information: Prediction or bimodal integration?

What happens if a visual cue misleads auditory expectations? Previous studies revealed an early visuo-auditory incongruency effect, so-called incongruency response (IR) of the auditory event-related brain potential (ERP), occurring 100 ms after onset of the sound being incongruent to the preceding visual cue. So far, this effect has been ascribed to reflect the mismatch between auditory sensory expectation activated by visual predictive information and the actual sensory input. Thus, an IR should be confined to an asynchronous presentation of visual cue and sound. Alternatively, one could argue that frequently presented congruent visual-cue-sound combinations are integrated into a bimodal representation whereby violation of the visual-auditory relationship results in a bimodal feature mismatch (the IR should be obtained with asynchronous and with synchronous presentation). In an asynchronous condition, an either high-pitched or low-pitched sound was preceded by a visual note symbol presented above or below a fixation cross (90% congruent; 10% incongruent), while in a synchronous condition, both were presented simultaneously. High-pitched and low-pitched sounds were presented with different probabilities (83% vs. 17%) to form a strong association between bimodal stimuli. In both conditions, tones with pitch incongruent with the location of the note symbols elicited incongruency effects in the N2 and P3 ERPs; however, the IR was only elicited in the asynchronous condition. This finding supports the sensorial prediction error hypothesis stating that the amplitude of the auditory ERP 100 ms after sound onset is enhanced in response to unexpected compared with expected but otherwise identical sounds.

What exactly is missing here? The sensory processing of unpredictable omissions is modulated by the specificity of expected action-effects.

We select our actions according to the desired outcomes; for instance, piano players press certain keys to generate specific musical notes. It is well-described that the omission of a predicted action-effect may elicit prediction error signals in the brain, but what happens in the case of simultaneous effector-specific (by contrast to effector-unspecific) predictions? To answer this question, we asked participants to press left and right keys to generate tones A and B; based on the action-effect association, the tones' identity was either predictable or unpredictable, while rarely, the expected input was omitted. Crucially, the data show that omissions following hand-specific associations reliably elicited a late omission N1 (oN1) component, by contrast to the hand-unspecific associations, where the late oN1 was rather weak. An additional condition where both key-presses generated a unique tone was implemented. Here, rare omissions of the expected tone generated both early and late oN1 responses, by contrast to the condition in which two simultaneous action-effect representations had to be maintained, where only late oN1 responses were elicited. Finally, omission P3 (oP3) responses were strongly elicited for all omission types without differences, indicating that a general expectation based on a tone presentation (rather than which tone), is likely indexed at this stage. The present results emphasize the top-down effects of action intention on the sensory processing of omissions, where unspecific (vs. specific) and multiple (vs. single) action-effect representations are associated with processing costs at the early sensory levels.

Interpretation bias and contamination-based obsessive-compulsive symptoms influence emotional intensity related to disgust and fear.

Biased processing of disgust-related stimuli is increasingly discussed in addition to fear-related processing as a maintenance factor for contamination-based obsessive-compulsive disorder (C-OCD). However, the differential impact of fear and disgust on biased processing in C-OCD is not yet completely understood. Because it is difficult to distinguish the two emotions in self-report assessment by directly addressing the specific emotions, a text paragraph-based interpretation bias paradigm was applied to more implicitly assess emotions. For the text-based interpretation bias paradigm, disgust-related, fear-related, disgust-fear-ambiguous and neutral text paragraphs describing everyday life situations were developed and validated in a pre-study (N = 205). Fifty-nine healthy participants watched either disgust- or fear-inducing movies and afterwards rated their experienced emotional response to the text paragraphs. The results show that fear and disgust components of an emotional response to mixed-emotional situations are strongly influenced by the situational context, and across the levels of trait contamination fear people did not differ in their fear experiences to everyday situations (which was overall strong), but in their disgust experiences. These findings highlight the strength of situational context on interpretation bias for mixed-emotional disorders and the important role of disgust for C-OCD.