Belief Updating during Social Interactions: Neural Dynamics and Causal Role of Dorsomedial Prefrontal Cortex.

In competitive interactions, humans have to flexibly update their beliefs about another person's intentions in order to adjust their own choice strategy, such as when believing that the other may exploit their cooperativeness. Here we investigate both the neural dynamics and the causal neural substrate of belief updating processes in humans. We used an adapted prisoner's dilemma game in which participants explicitly predicted the coplayer's actions, which allowed us to quantify the prediction error between expected and actual behavior. First, in an EEG experiment, we found a stronger medial frontal negativity (MFN) for negative than positive prediction errors, suggesting that this medial frontal ERP component may encode unexpected defection of the coplayer. The MFN also predicted subsequent belief updating after negative prediction errors. In a second experiment, we used transcranial magnetic stimulation (TMS) to investigate whether the dorsomedial prefrontal cortex (dmPFC) causally implements belief updating after unexpected outcomes. Our results show that dmPFC TMS impaired belief updating and strategic behavioral adjustments after negative prediction errors. Taken together, our findings reveal the time course of the use of prediction errors in social decisions and suggest that the dmPFC plays a crucial role in updating mental representations of others' intentions.

My choice, my actions: self-determination, not instrumental value of outcomes enhances outcome monitoring during learning.

Freedom of choice enhances our sense of agency. During goal-directed behavior, the freedom to choose between different response options increases the neural processing of positive and negative feedback, indicating enhanced outcome monitoring under conditions of high agency experience. However, it is unclear whether this enhancement is predominantly driven by an increased salience of self- compared to externally determined action outcomes or whether differences in the perceived instrumental value of outcomes contribute to outcome monitoring in goal-directed tasks. To test this, we recorded electroencephalography while participants performed a reinforcement learning task involving free choices, action-relevant forced choices, and action-irrelevant forced choices. We observed larger midfrontal theta power and N100 amplitudes for feedback following free choices compared with action-relevant and action-irrelevant forced choices. In addition, a Reward Positivity was only present for free but not forced choice outcomes. Crucially, our results indicate that enhanced outcome processing is not driven by the relevance of outcomes for future actions but rather stems from the association of outcomes with recent self-determined choice. Our findings highlight the pivotal role of self-determination in tracking the consequences of our actions and contribute to an understanding of the cognitive processes underlying the choice-induced facilitation in outcome monitoring.

Listen to your heart: Trade-off between cardiac interoceptive processing and visual exteroceptive processing.

Internal bodily signals, such as heartbeats, can influence conscious perception of external sensory information. Spontaneous shifts of attention between interoception and exteroception have been proposed as the underlying mechanism, but direct evidence is lacking. Here, we used steady-state visual evoked potential (SSVEP) frequency tagging to independently measure the neural processing of visual stimuli that were concurrently presented but varied in heartbeat coupling in healthy participants. Although heartbeat coupling was irrelevant to participants' task of detecting brief color changes, we found decreased SSVEPs for systole-coupled stimuli and increased SSVEPs for diastole-coupled stimuli, compared to non-coupled stimuli. These results suggest that attentional and representational resources allocated to visual stimuli vary according to fluctuations in cardiac-related signals across the cardiac cycle, reflecting spontaneous and immediate competition between cardiac-related signals and visual events. Furthermore, frequent coupling of visual stimuli with stronger cardiac-related signals not only led to a larger heartbeat evoked potential (HEP) but also resulted in a smaller color change evoked N2 component, with the increase in HEP amplitude associated with a decrease in N2 amplitude. These findings indicate an overall or longer-term increase in brain resources allocated to the internal domain at the expense of reduced resources available for the external domain. Our study highlights the dynamic reallocation of limited processing resources across the internal-external axis and supports the trade-off between interoception and exteroception.

Prepared to stop: how sense of agency in a preceding trial modulates inhibitory control in the current trial.

Feeling in control of actions and events can enhance motivation for further actions. How this sense of agency (SoA) in fact influences flexible motor control remains poorly understood. Here, we investigated the effect of SoA on subsequent response inhibition in a modified go/no-go task with EEG recordings. We manipulated participants' SoA by varying the presence, predictability, and emotional valence of a visual outcome for a given motor action. When participants unexpectedly did not receive any visible outcome following their action on trial n - 1, they exhibited slower responses and lower hit rates to the go signal but higher rates of successful inhibition to the no-go signal on trial n, regardless of the emotional valence of the expected action outcome. Furthermore, enhanced inhibitory tendencies were accompanied by reduced N2 and P3 amplitudes, midfrontal theta power, and theta synchronization between midfrontal and medial to parietal areas, indicating that less top-down control is required for successful response inhibition on trial n after experiencing low SoA on trial n - 1. These findings suggest that feeling less in control in a preceding trial makes it easier to implement inhibitory control in the current trial, thereby providing new insights into the role of SoA in goal-directed behavior.

Function without feeling: neural reactivity and intercommunication during flexible motor adjustments evoked by emotional and neutral stimuli.

Motor conflicts arise when we need to quickly overwrite prepotent behavior. It has been proposed that affective stimuli modulate the neural processing of motor conflicts. However, previous studies have come to inconsistent conclusions regarding the neural impact of affective information on conflict processing. We employed functional magnetic resonance imaging during a Go/Change-Go task, where motor conflicts were either evoked by neutral or emotionally negative stimuli. Dynamic causal modeling was used to investigate how motor conflicts modulate the intercommunication between the anterior cingulate cortex (ACC) and the anterior insula (AI) as 2 central regions for cognitive control. Conflicts compared to standard actions were associated with increased BOLD activation in several brain areas, including the dorsal ACC and anterior insula. There were no differences in neural activity between emotional and non-emotional conflict stimuli. Conflicts compared to standard actions lowered neural self-inhibition of the ACC and AI and led to increased effective connectivity from the ACC to AI contralateral to the acting hand. Thus, our study indicates that neural conflict processing is primarily driven by the functional relevance of action-related stimuli, not their inherent affective meaning. Furthermore, it sheds light on the role of interconnectivity between ACC and AI for the implementation of flexible behavioral change.

Flexible Changes in Attentional Focus and Task Rules Rely on A Shared Set of Frontoparietal Oscillatory Dynamics.

Flexible changes in behavior can involve changes in the processing of external information (i.e., shifts in attention between different stimuli) or internal information (i.e., shifts in task rules stored in memory). However, it is unclear if different types of flexible change rely on separate, domain-specific neural processes or on a domain-general system, which enables flexible actions independent of the type of change needed. In the current study, participants performed a task switching procedure while we measured neural oscillations via EEG. Importantly, we independently manipulated the need to switch attention between 2 types of stimuli, as well as the need to switch between two sets of stimuli-response rules stored in memory. Both attentional and rule switches significantly increased error rates and RTs. On a neural level, both types of changes were associated with a widespread decrease in alpha power, predominantly over the parietal cortex. Attentional switches and rule switches showed a subadditive interaction effect on both participants' performance as well as on their alpha power reactivity. This indicates that implementing both changes at the same time was more efficient than implementing each individual change separately. Independent of the presence or absence of either attentional or rule switches, higher frontal theta power and lower parietal/posterior alpha power predicted faster responses on correct trials. Our study suggests that flexible behavior relies on domain-general frontal and parietal oscillatory dynamics, which enable efficient implementation of goal-directed actions independent of which aspects of the task change.

Motor Interference, But Not Sensory Interference, Increases Midfrontal Theta Activity and Brain Synchronization during Reactive Control.

Cognitive control helps us to overcome task interference in challenging situations. Resolving conflicts because of interfering influences is believed to rely on midfrontal theta oscillations. However, different sources of interference necessitate different types of control. Attentional control is needed to suppress salient distractors. Motor control is needed to suppress goal-incompatible action impulses. While previous studies mostly studied the additive effects of attentional and motor conflicts, we independently manipulated the need for attentional control (via visual distractors) and motor control (via unexpected response deviations) in an EEG study with male and female humans. We sought to find out whether these different types of control rely on the same midfrontal oscillatory mechanisms. Motor conflicts, but not attentional conflicts, elicited increases in midfrontal theta power during conflict resolution. Independent of the type of conflict, theta power was predictive of motor slowing. Connectivity analysis via phase-based synchronization indicated a widespread increase interbrain connectivity for motor conflicts, but a midfrontal-to-posterior decrease in connectivity for attentional conflicts. For each condition, we found stronger midfrontal connectivity with the parietal region contralateral to, rather than ipsilateral to, the acting hand. Parietal lateralization in connectivity was strongest for motor conflicts. Previous studies suggested that midfrontal theta oscillations might represent a general control mechanism, which aids conflict resolution independent of the conflict domain. In contrast, our results show that oscillatory theta dynamics during reactive control mostly reflect motor-related adjustments.SIGNIFICANCE STATEMENT Humans need to exercise self-control over both their attention (to avoid distraction) and their motor activity (to suppress inappropriate action impulses). Midfrontal theta oscillations have been assumed to indicate a general control mechanism, which help to exert top-down control during both motor and sensory interference. We are using a novel approach for the independent manipulation of attentional and motor control to show that increases in midfrontal theta power and brainwide connectivity are linked to the top-down adjustments of motor responses, not sensory interference. These findings clarify the function of midfrontal theta dynamics as a key aspect of neural top-down control and help to dissociate domain-general from motor-specific aspects of self-control.

Preparing for Success: Neural Frontal Theta and Posterior Alpha Dynamics during Action Preparation Predict Flexible Resolution of Cognitive Conflicts.

Cognitive conflicts typically arise in situations that call for sudden changes in our behavior. Resolving cognitive conflicts is challenging and prone to errors. Humans can improve their chances to successfully resolve conflicts by mentally preparing for potential behavioral adjustments. Previous studies indicated that neural theta oscillations (4-7 Hz), as well as alpha oscillations (8-14 Hz), are reflective of cognitive control processes during conflict resolution. However, the role or neural oscillations for conflict preparation is still unclear. Therefore, the aim of the current study was to determine which oscillatory changes during conflict preparation predict subsequent resolution success. Participants performed a cued change-signal task, in which an anticipatory cue indicated if the upcoming trial might contain a cognitive conflict or not. Oscillatory activity was assessed via EEG. Cues that indicated that a conflict might arise compared with cues that indicated no conflict led to increases, directly followed by decreases, in theta power, as well as to decreases in alpha power. These cue-induced changes in theta and alpha oscillations occurred widespread across the cortex. Importantly, successful compared with failed conflict trials were characterized by selective increases in frontal theta power, as well as decreases in posterior alpha power during preparation. In addition, higher frontal theta power and lower posterior alpha power during preparation predicted faster conflict resolution. Our study shows that increases in frontal theta power, as well as decreases in posterior alpha power, are markers of optimal preparation for situations that necessitate flexible changes in behavior.

Multisensory integration of anticipated cardiac signals with visual targets affects their detection among multiple visual stimuli.

Many studies have elucidated the multisensory processing of different exteroceptive signals (e.g., auditory-visual stimuli), but less is known about the multisensory integration of interoceptive signals with exteroceptive information. Here, we investigated the perceptual outcomes and electrophysiological brain mechanisms of cardio-visual integration by using participants' electrocardiogram signals to control the color change of a visual target in dynamically changing displays. Reaction times increased when the target change coincided with strong cardiac signals concerning the state of cardiovascular arousal (i.e., presented at the end of ventricular systole), compared to when the target change occurred at a time when cardiac arousal was relatively low (i.e., presented at the end of ventricular diastole). Moreover, the concurrence of the target change and cardiac arousal signals modulated the event-related potentials and the beta power in an early period (~100 ms after stimulus onset), and decreased the N2pc and the beta lateralization in a later period (~200 ms after stimulus onset). Our results suggest that the multisensory integration of anticipated cardiac signals with a visual target negatively affects its detection among multiple visual stimuli, potentially by suppressing sensory processing and reducing attention toward the visual target. This finding highlights the role of cardiac information in visual processing and furthers our understanding of the brain dynamics underlying multisensory perception involving both interoception and exteroception.

From the inside out: Interoceptive feedback facilitates the integration of visceral signals for efficient sensory processing.

Neuroscientific studies have mainly focused on the way humans perceive and interact with the external world. Recent work in the interoceptive domain indicates that the brain predictively models information from inside the body such as the heartbeat and that the efficiency with which this is executed can have implications for exteroceptive processing. However, to date direct evidence underpinning these hypotheses is lacking. Here, we show how the brain predictively refines neural resources to process afferent cardiac feedback and uses these interoceptive cues to enable more efficient processing of external sensory information. Participants completed a repetition-suppression paradigm consisting of a neutral repeating face. During the first face presentation, they heard auditory feedback of their heartbeat which either coincided with the systole of the cardiac cycle, the time at which cardiac events are registered by the brain or the diastole during which the brain receives no internal cardiac feedback. We used electroencephalography to measure the heartbeat evoked potential (HEP) as well as auditory (AEP) and visual evoked potentials (VEP). Exteroceptive cardiac feedback which coincided with the systole produced significantly higher HEP amplitudes relative to feedback timed to the diastole. Elevation of the HEP in this condition was followed by significant suppression of the VEP in response to the repeated neutral face and a stepwise decrease of AEP amplitude to repeated heartbeat feedback. Our results hereby show that exteroceptive heartbeat feedback coinciding with interoceptive signals at systole enhanced interoceptive cardiac processing. Furthermore, the same cue facilitating interoceptive integration enabled efficient suppression of a visual stimulus, as well as repetition suppression of the AEP across successive auditory heartbeat feedback. Our findings provide evidence that the alignment of external to internal signals can enhance the efficiency of interoceptive processing and that cues facilitating this process in either domain have beneficial effects for internal as well as external sensory processing.

Learning something new versus changing your ways: Distinct effects on midfrontal oscillations and cardiac activity for learning and flexible adjustments.

We need to be able to learn new behaviour, but also be capable of changing existing routines, when they start conflicting with our long-term goals. Little is known about to what extent blank-slate learning of new and adjustment of existing behavioural routines rely on different neural and bodily mechanisms. In the current study, participants first acquired novel stimulus-response contingencies, which were subsequently randomly changed to create the need for flexible adjustments. We measured midfrontal theta oscillations via EEG as an indicator of neural conflict processing, as well as heart rate as a proxy of autonomic activity. Participants' trial-wise learning progress was estimated via computation modelling. Theta power and heart rate significantly differed between correct and incorrect trials. Differences between correct and incorrect trials in both neural and cardiac feedback processing were more pronounced for adjustments compared to blank-slate learning. This indicates that both midfrontal and cardiac processing are sensitive to changes in stimulus-response contingencies. Increases in individual learning rates predicted lower impact of performance feedback on midfrontal theta power, but higher impact on heart rate. This suggests that cardiac and midfrontal reactivity are partially reflective of different mechanisms related to feedback learning. Our results shed new light on the role of neural and autonomic mechanisms for learning and behavioural adjustments.

I feel what I do: Relating interoceptive processes and reward-related behavior.

Interoceptive signalling has been shown to contribute to action regulation and action experience. Here, we assess whether motor behaviour can be influenced by anticipated homeostatic feeling states induced through different predictable contexts. Participants performed a reward incentive paradigm in which accurate responses increased (gain) or avoided the depletion (averted loss) of a credit score. Across two types of blocks, we varied the predictability of the outcome state. In predictable blocks, a cue signaled a gain, loss or control trial (motor response did not affect the credit score). This allowed participants to anticipate the interoceptive feeling state associated with the outcome. In unpredictable blocks, the cue had no relation to the type of outcome. Thus, participants were unable to anticipate the feeling state it produced. Via EEG, we measured the Heartbeat Evoked Potential (HEP) and the Contingent Negative Variation (CNV) as indices of interoceptive and motor processing respectively. In addition, we measured feedback P3 amplitude following outcome presentation and accuracy and reaction times of the required motor response. We observed higher HEP and CNV amplitudes as well as faster and more accurate motor responses in predictable compared to unpredictable outcome blocks. Similarly, feedback-related P3 amplitudes were significantly lower for predictable relative to unpredictable outcomes. Crucially, HEP amplitudes measured prior to feedback predicted feedback-related P3 amplitudes for anticipated outcome events. Results suggest that accurate anticipation of homeostatic feeling states associated with gain, loss or control outcomes facilitates motor execution and outcome evaluation. Findings are hereby the first to empirically assess the link between interoceptive and motor domains and provide primary evidence for a joint processing structure.

Affective interoceptive inference: Evidence from heart-beat evoked brain potentials.

The perception of internal bodily signals (interoception) is central to many theories of emotion and embodied cognition. According to recent theoretical views, the sensory processing of visceral signals such as one's own heartbeat is determined by top-down predictions about the expected interoceptive state of the body (interoceptive inference). In this EEG study we examined neural responses to heartbeats following expected and unexpected emotional stimuli. We used a modified stimulus repetition task in which pairs of facial expressions were presented with repeating or alternating emotional content, and we manipulated the emotional valence and the likelihood of stimulus repetition. We found that affective predictions of external socially relevant information modulated the heartbeat-evoked potential, a marker of cardiac interoception. Crucially, the HEP changes highly relied on the expected emotional content of the facial expression. Thus, expected negative faces led to a decreased HEP amplitude, whereas such an effect was not observed after an expected neutral face. These results suggest that valence-specific affective predictions, and their uniquely associated predicted bodily sensory state, can reduce or amplify cardiac interoceptive responses. In addition, the affective repetition effects were dependent on repetition probability, highlighting the influence of top-down exteroceptive predictions on interoception. Our results are in line with recent models of interoception supporting the idea that predicted bodily states influence sensory processing of salient external information.

Sensory attenuation of self-produced signals does not rely on self-specific motor predictions.

Sensory events produced by ourselves are known to lead to lower neural and perceptual impact than sensory events from other environmental sources. This sensory attenuation is widely assumed to result from control processes that are specific to our own motor actions, potentially helping us to distinguish effects produced by ourselves and others. However, previous research cannot rule out that the putative self-attenuation in fact reflect actor-independent, general predictive mechanisms, which, in direct comparison, just highlight external events due to lower predictability of their onset and thus higher surprise. By measuring the auditory-evoked N1 component, we show that self-generation of sounds only lead to cortical attenuation when the onset of other-generated sounds is less predictable due to the absence of any predictive cues. The presence of a cue predicting the onset of auditory stimuli, in contrast, led to a reversal of the attenuation effect, with lower N1 amplitudes for other-generated sounds in contrast to self-generated sounds. Thus, contrary to prevalent assumptions sensory attenuation is not bound to self-generation per se. Rather, it appears to be the result of general mechanisms that does not reliably and selectively attenuate self-induced stimulation but is determined by a flexible processing of sensory input based on its predictability, contextual relevance and attentional salience.

Imitating others' actions: transcranial magnetic stimulation of the parietal opercula reveals the processes underlying automatic imitation.

The human tendency to imitate gestures performed by conspecifics is automatic in nature. However, whether this automatic imitation can be considered as a true imitative phenomenon or only as a special instance of spatial compatibility is still being debated. New evidence suggests that automatic imitation, otherwise known as 'imitative compatibility', shall be considered as a phenomenon that operates independently from spatial compatibility. So far there are only a few investigations directly aimed at identifying the neural structures dedicated to this process. In the present study, we applied double-pulse transcranial magnetic stimulation (TMS) over the parietal opercula to further investigate the role of these regions in coding imitative compatibility. We found that a temporary disruption of parietal opercula caused the reduction of the imitative compatibility relative to the sham condition. In particular, the TMS interference with the parietal opercula's activity modulated the imitative compatibility but not the spatial compatibility, suggesting that these two processes are likely to be independent.

Judging roughness by sight--a 7-Tesla fMRI study on responsivity of the primary somatosensory cortex during observed touch of self and others.

Observing another person being touched activates our own somatosensory system. Whether the primary somatosensory cortex (S1) is also activated during the observation of passive touch, and which subregions of S1 are responsible for self- and other-related observed touch is currently unclear. In our study, we first aimed to clarify whether observing passive touch without any action component can robustly increase activity in S1. Secondly, we investigated whether S1 activity only increases when touch of others is observed, or also when touch of one's own body is observed. We were particularly interested in which subregions of S1 are responsible for either process. We used functional magnetic resonance imaging at 7 Tesla to measure S1 activity changes when participants observed videos of their own or another's hand in either egocentric or allocentric perspective being touched by different pieces of sandpaper. Participants were required to judge the roughness of the different sandpaper surfaces. Our results clearly show that S1 activity does increase in response to observing passive touch, and that activity changes are localized in posterior but not in anterior parts of S1. Importantly, activity increases in S1 were particularly related to observing another person being touched. Self-related observed touch, in contrast, caused no significant activity changes within S1. We therefore assume that posterior but not anterior S1 is part of a system for sharing tactile experiences with others.

The influence of visual training on predicting complex action sequences.

Linking observed and executable actions appears to be achieved by an action observation network (AON), comprising parietal, premotor, and occipitotemporal cortical regions of the human brain. AON engagement during action observation is thought to aid in effortless, efficient prediction of ongoing movements to support action understanding. Here, we investigate how the AON responds when observing and predicting actions we cannot readily reproduce before and after visual training. During pre- and posttraining neuroimaging sessions, participants watched gymnasts and wind-up toys moving behind an occluder and pressed a button when they expected each agent to reappear. Between scanning sessions, participants visually trained to predict when a subset of stimuli would reappear. Posttraining scanning revealed activation of inferior parietal, superior temporal, and cerebellar cortices when predicting occluded actions compared to perceiving them. Greater activity emerged when predicting untrained compared to trained sequences in occipitotemporal cortices and to a lesser degree, premotor cortices. The occipitotemporal responses when predicting untrained agents showed further specialization, with greater responses within body-processing regions when predicting gymnasts' movements and in object-selective cortex when predicting toys' movements. The results suggest that (1) select portions of the AON are recruited to predict the complex movements not easily mapped onto the observer's body and (2) greater recruitment of these AON regions supports prediction of less familiar sequences. We suggest that the findings inform both the premotor model of action prediction and the predictive coding account of AON function.

Musical groove modulates motor cortex excitability: a TMS investigation.

Groove is often described as a musical quality that can induce movement in a listener. This study examines the effects of listening to groove music on corticospinal excitability. Musicians and non-musicians listened to high-groove music, low-groove music, and spectrally matched noise, while receiving single-pulse transcranial magnetic stimulation (TMS) over the primary motor cortex either on-beat or off-beat. We examined changes in the amplitude of the motor-evoked potentials (MEPs), recorded from hand and arm muscles, as an index of activity within the motor system. Musicians and non-musicians rated groove similarly. MEP results showed that high-groove music modulated corticospinal excitability, whereas no difference occurred between low-groove music and noise. More specifically, musicians' MEPs were larger with high-groove than low-groove music, and this effect was especially pronounced for on-beat compared to off-beat pulses. These results indicate that high-groove music increasingly engages the motor system, and the temporal modulation of corticospinal excitability with the beat could stem from tight auditory-motor links in musicians. Conversely, non-musicians' MEPs were smaller for high-groove than low-groove music, and there was no effect of on- versus off-beat pulses, potentially stemming from suppression of overt movement. In sum, high-groove music engages the motor system, and previous training modulates how listening to music with a strong groove activates the motor system.

Distinguishing self and other in joint action. Evidence from a musical paradigm.

The capacity to distinguish between one's own and others' behavior is a cognitive prerequisite for successful joint action. We employed a musical joint action task to investigate how the brain achieves this distinction. Pianists performed the right-hand part of piano pieces, previously learned bimanually, while the complementary left-hand part either was not executed or was (believed to be) played by a co-performer. This experimental setting served to induce a co-representation of the left-hand part reflecting either the self or the co-performer. Single-pulse transcranial magnetic stimulation was applied to the right primary motor cortex and motor-evoked potentials (MEPs) were recorded from the resting left forearm. Results show that corticospinal excitability was modulated by whether the representation of the left hand was associated with the self or the other, with the MEP amplitude being low and high, respectively. This result remained unchanged in a separate session where participants could neither see nor hear the other but still infer his presence by means of contextual information. Furthermore, the amplitude of MEPs associated with co-performer presence increased with pianists' self-reported empathy. Thus, the sociality of the context modulates action attribution at the level of the motor control system.

Freedom of choice boosts midfrontal theta power during affective feedback processing of goal-directed actions.

Sense of agency, the feeling of being in control of one's actions and their effects, is particularly relevant during goal-directed actions. During feedback learning, action effects provide information about the best course of action to reinforce positive and prevent negative outcomes. However, it is unclear whether agency experience selectively affects the processing of negative or positive feedback during the performance of goal-directed actions. As an important marker of feedback processing, we examined agency-related changes in midfrontal oscillatory activity in response to performance feedback using electroencephalography. Thirty-three participants completed a reinforcement learning task during which they received positive (monetary gain) or negative (monetary loss) feedback following item choices made either by themselves (free-choice) or by the computer (forced-choice). Independent of choice context, midfrontal theta activity was more enhanced for negative than positive feedback. In addition, free, compared to forced choices increased midfrontal theta power for both gain and loss feedback. These results indicate that freedom of choice in a motivationally salient learning task leads to a general enhancement in the processing of affective action outcomes. Our findings contribute to an understanding of the neuronal mechanisms underlying agency-related changes during action regulation and indicate midfrontal theta activity as a neurophysiological marker important for the monitoring of affective action outcomes, irrespective of feedback valence.