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.

Ready to go: Higher sense of agency enhances action readiness and reduces response inhibition.

Sense of agency is the subjective feeling of being in control of one's actions and their effects. Many studies have elucidated the cognitive and sensorimotor processes that drive this experience. However, less is known about how sense of agency influences flexible cognitive and motor control. Here, we investigated the effect of sense of agency on subsequent action regulation using a modified Go/No-Go task. In Experiment 1, we modulated participants' sense of agency by varying the occurrence of action outcomes (present vs. absent) both locally on a trial-by-trial basis and globally in terms of the overall probability of action outcomes within a block of trials (high vs. low). Importantly, we investigated how this manipulation influenced participants' responses to subsequent Go, No-Go, or Free-Choice cues. When participants' previous action led to an outcome (i.e., a happy face) compared with no outcome, they responded more accurately and faster to Go cues, reacted less accurately to No-Go cues, as well as made go decisions more frequently and faster to Free-Choice cues. These effects were even stronger when action outcomes occurred more frequently overall in a given block or in several previous trials. Experiment 2 further demonstrated that the effects of action outcome manipulation on subsequent action regulation were independent of the emotional valence of the action outcome (i.e., a happy or an angry face). Our results suggest that a higher sense of agency as induced by the presence of action outcomes enhanced action readiness and suppressed response inhibition. These findings highlight the impact of the control felt on the control used in action regulation, thereby providing new insights into the functional significance of the sense of agency on human 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.

Clinical Manifestations of Body Memories: The Impact of Past Bodily Experiences on Mental Health.

Bodily experiences such as the feeling of touch, pain or inner signals of the body are deeply emotional and activate brain networks that mediate their perception and higher-order processing. While the ad hoc perception of bodily signals and their influence on behavior is empirically well studied, there is a knowledge gap on how we store and retrieve bodily experiences that we perceived in the past, and how this influences our everyday life. Here, we explore the hypothesis that negative body memories, that is, negative bodily experiences of the past that are stored in memory and influence behavior, contribute to the development of somatic manifestations of mental health problems including somatic symptoms, traumatic re-experiences or dissociative symptoms. By combining knowledge from the areas of cognitive neuroscience and clinical neuroscience with insights from psychotherapy, we identify Clinical Body Memory (CBM) mechanisms that specify how mental health problems could be driven by corporeal experiences stored in memory. The major argument is that the investigation of the neuronal mechanisms that underlie the storage and retrieval of body memories provides us with empirical access to reduce the negative impact of body memories on mental health.

The Interplay Between Affective Processing and Sense of Agency During Action Regulation: A Review.

Sense of agency is the feeling of being in control of one's actions and their perceivable effects. Most previous research identified cognitive or sensory determinants of agency experience. However, it has been proposed that sense of agency is also bound to the processing of affective information. For example, during goal-directed actions or instrumental learning we often rely on positive feedback (e.g., rewards) or negative feedback (e.g., error messages) to determine our level of control over the current task. Nevertheless, we still lack a scientific model which adequately explains the relation between affective processing and sense of agency. In this article, we review current empirical findings on how affective information modulates agency experience, and, conversely, how sense of agency changes the processing of affective action outcomes. Furthermore, we discuss in how far agency-related changes in affective processing might influence the ability to enact cognitive control and action regulation during goal-directed behavior. A preliminary model is presented for describing the interplay between sense of agency, affective processing, and action regulation. We propose that affective processing could play a role in mediating the influence between subjective sense of agency and the objective ability to regulate one's behavior. Thus, determining the interrelation between affective processing and sense of agency will help us to understand the potential mechanistic basis of agency experience, as well as its functional significance for goal-directed behavior.

Action inhibition in individuals with high obsessive-compulsive trait of incompleteness: An ERP study.

BACKGROUND: Missing action completion signals are assumed to trigger repetitive behavior and feelings of the action "not being right". This proposal is based mostly on individual's self-reports. Here, we investigated the influence of experimentally manipulated action completion experience and the obsessive-compulsive (OC) trait of incompleteness on behavioral and neurophysiological measures of action inhibition. METHODS: Action completion was manipulated in an adapted Go/NoGo task, and OC trait incompleteness was assessed in healthy participants. More commission errors and faster responses were expected after missing action completion, especially for individuals with high OC trait incompleteness. The inhibition-related event-related potentials (ERPs) N200 and P300 were also measured. RESULTS: High OC trait incompleteness led to more errors following omitted- and faster responses during commission errors following incongruent outcomes. Furthermore, lower N200 was associated with worse response inhibition, and high OC trait incompleteness was associated with reduced N200, but not reduced P300 amplitude. These findings provide evidence that trait-like feelings of incompleteness may underlie maladaptive action repetition and impaired inhibitory control as observed in OCD.

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.

Interoceptive cardiac expectations to emotional stimuli predict visual perception.

Previous work exploring the interplay between the processing of internal bodily signals and the perception of external events suggests a correlative relationship between both domains. Here, we demonstrate an experimentally induced link between visual perception and the emotional modulation of interoceptive cardiac signals. We pair an emotion repetition-suppression paradigm shown to modulate neural responses to heartbeats with a previously reported visual detection task. Using electrophysiological and cardiac measurements, we show that, relative to alternating facial expressions of emotions, repeating facial expressions evoke stable patterns of cardiac responses and that this iteration of the cardiac response leads to a distinct modulation of the heartbeat evoked potential (HEP) amplitude. Crucially, we were able to modulate the direction of the amplitude change to increase HEP expression in response to repeated emotionally neutral faces and decrease it by repeating angry faces. Visual detection accuracy was significantly higher in repeating, relative to alternating face trials. Furthermore, reduced HEP amplitude to repeated angry faces acted as a direct predictor of subsequently elevated detection performance. Our findings demonstrate that accurate top-down anticipation of the heartbeat signal coincides with enhanced exteroceptive visual perception, particularly in the context of emotional events. Findings hereby highlight the integrated nature of internal and external stimulus processing and implicate predictive interoceptive processes induced by emotions for the conscious perception of sensory input activated from outside of the body. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

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.

Sensitivity to CT-optimal, Affective Touch Depends on Adult Attachment Style.

Affective touch supports affiliative bonds and social cognition. In particular, gentle, stroking touch, which has recently been associated with the C Tactile (CT) system, is typically perceived as pleasant and prosocial. However, it remains unknown whether pre-existing models of social relating influence the perception of CT-optimal touch. In this study (N = 44 adults), we examined how individual differences in attachment styles relate to the perception of CT-optimal touch, as well as to a different modality of interoception, namely heartbeat perception. Using the gold-standard assessment of attachment (Adult Attachment Interview), we found that insecure attachment was associated with reduced pleasantness discrimination between CT-optimal vs. non-CT optimal touch. Acknowledging the different traditions in measuring attachment, we also used a well-validated self-report questionnaire that pertains to explicit representations of current close relationships. Using this measure, we found that higher scores in attachment anxiety (but not attachment avoidance) were associated with reduced pleasantness discrimination between CT-optimal vs. non-CT optimal touch. Attachment patterns (in both measures) were not related to cardiac perception accuracy. These results corroborate and extend previous literature on CT-optimal touch and its relation with affiliative bonds and social cognition. Given that attachment was not related to perceived cardiac accuracy, these findings point to the specificity of the relationship between CT-optimal touch and attachment.

Dissociable sources of erogeneity in social touch: Imagining and perceiving C-Tactile optimal touch in erogenous zones.

Previous research points to two major hypotheses regarding the mechanisms by which touch can be experienced as erotogenic. The first concerns the body part to which touch is applied (erogenous zones) and the second the modality of touch (sensual touch optimal in activating C Tactile afferents). In this study, we explored for the first time the relation between those two mechanisms in actual and imagined social touch. In a first experiment, we randomly assigned "Giver" and "Receiver" roles within 19 romantic couples (20 females, 18 males, age 32.34 ± 8.71SD years) and asked the "Giver" to apply CT-optimal (3 cm/s) vs. CT-suboptimal (18 cm/s) touch on an erogenous (neck) vs. non-erogenous zone (forehead) of their partner. We then obtained ratings of pleasantness and sexual arousal from both "Receivers" and "Givers". In a second experiment, 32 healthy females (age 25.16 ± 5.91SD years) were asked to imagine CT-optimal vs. CT-suboptimal stimulation (stroking vs. patting) and velocity (3 cm/s vs. 18 cm/s) on different erogenous vs. non-erogenous zones and rate pleasantness. While both erogenous body part and CT-optimal, sensual touch were found to increase pleasant and erotic sensations, the results showed a lack of an interaction. Furthermore, pleasantness was induced by mere imagination of touch without any tactile stimulation, and touch that was sexually arousing for the receiver was rated as more sexually arousing for the giver as well, pointing to top-down, learned expectations of sensory pleasure and erogeneity. Taken together, these studies provide the first direct evidence that while both the body location to which touch is applied and the mode of touch contribute to pleasant and erotic sensations, these two factors appear to mediate subjective pleasantness and erogeneity by, at least partly, independent mechanisms.

Cardiac interoceptive learning is modulated by emotional valence perceived from facial expressions.

Interoception refers to the processing of homeostatic bodily signals. Research demonstrates that interoceptive markers can be modulated via exteroceptive stimuli and suggests that the emotional content of this information may produce distinct interoceptive outcomes. Here, we explored the impact of differently valenced exteroceptive information on the processing of interoceptive signals. Participants completed a repetition-suppression paradigm viewing repeating or alternating faces. In experiment 1, faces wore either angry or pained expressions to explore the interoceptive response to different types of negative stimuli in the observer. In experiment 2, expressions were happy or sad to compare interoceptive processing of positive and negative information. We measured the heartbeat evoked potential (HEP) and visual evoked potentials (VEPs) as a respective marker of intero- and exteroceptive processing. We observed increased HEP amplitude to repeated sad and pained faces coupled with reduced HEP and VEP amplitude to repeated angry faces. No effects were observed for positive faces. However, we found a significant correlation between suppression of the HEP and VEP to repeating angry faces. Results highlight an effect of emotional expression on interoception and suggest an attentional trade-off between internal and external processing domains as a potential account of this phenomenon.

The Interaction between Interoceptive and Action States within a Framework of Predictive Coding.

The notion of predictive coding assumes that perception is an iterative process between prior knowledge and sensory feedback. To date, this perspective has been primarily applied to exteroceptive perception as well as action and its associated phenomenological experiences such as agency. More recently, this predictive, inferential framework has been theoretically extended to interoception. This idea postulates that subjective feeling states are generated by top-down inferences made about internal and external causes of interoceptive afferents. While the processing of motor signals for action control and the emergence of selfhood have been studied extensively, the contributions of interoceptive input and especially the potential interaction of motor and interoceptive signals remain largely unaddressed. Here, we argue for a specific functional relation between motor and interoceptive awareness. Specifically, we implicate interoceptive predictions in the generation of subjective motor-related feeling states. Furthermore, we propose a distinction between reflexive and pre-reflexive modes of agentic action control and suggest that interoceptive input may affect each differently. Finally, we advocate the necessity of continuous interoceptive input for conscious forms of agentic action control. We conclude by discussing further research contributions that would allow for a fuller understanding of the interaction between agency and interoceptive awareness.

Exteroceptive expectations modulate interoceptive processing: repetition-suppression effects for visual and heartbeat evoked potentials.

Interoception refers to the signaling of internal bodily commands. Here, we explore repetition suppression of intero- and exteroceptive neural markers to test whether the perception and predictability of exteroceptive stimulus material affects their expression. Participants completed a repetition suppression paradigm in which angry or neutral facial expressions repeated or alternated. Participants received either an implicit (experiment 1) or explicit (experiment 2) cue enabling the formation of expectations regarding the upcoming facial expression. We measured the heartbeat-evoked potential (HEP) indexing cardiac processing and visual evoked potentials (VEP) in response to viewing the second (repeated or alternated) face. Repeating angry facial expressions produced repetition suppression of both HEP and VEP amplitude while repeating neutral expressions led to repetition enhancement of HEP amplitude. This effect was magnified when participants were explicitly aware of predictive cues. Furthermore, repetition suppression of HEP amplitude correlated with neural attenuation of VEP activity. Results highlight repetition effects for interoceptive as well as exteroceptive neural markers and support top-down, expectation-based accounts of the phenomenon. Furthermore, results demonstrate that the perception of exteroceptive stimulus information has an effect on the processing of interoceptive signals and suggest a direct neural connection between the processing of external and internal sensory information.

Towards a common framework of grounded action cognition: Relating motor control, perception and cognition.

The relation between motor control and action cognition - including action-related thoughts and action-related perception - has been subject to controversial discussions in the last three decades. During these decades, cognitive neuroscience has been increasingly confronted with a huge variety of different accounts trying to understand and explain the relation between these systems, their interdependencies and the mediating mechanisms by establishing notions such as "internal models", "simulation" or "shared representation". These accounts, however, include a large array of partly overlapping, partly contradictory theories using similar terms for different mechanisms and different terms for similar mechanisms. In the absence of a systematic work-up and comparison, this array of accounts and theories leads to confusion in the field, duplication of experimental work, and unconnected parallelism of theory formation within and between different disciplines. Here we provide a systematic comparison of current models and prospective theories that deal with the relation between cognition, perception and motor control mechanisms. In a second step, we propose "grounded action cognition" as a comprehensive metatheoretical framework which defines different hypothetical possibilities of the relations between these domains, offers systematic insights into current models and theories and last but not least may help to increase comparability of empirical research in the domain of action and action cognition.

Active Interpersonal Touch Gives Rise to the Social Softness Illusion.

Social touch plays a powerful role in human life, with important physical and mental health benefits in development and adulthood. Touch is central in building the foundations of social interaction, attachment, and cognition, and early, social touch has unique, beneficial neurophysiological and epigenetic effects. The recent discovery of a separate neurophysiological system for affectively laden touch in humans has further kindled scientific interest in the area. Remarkably, however, little is known about what motivates and sustains the human tendency to touch others in a pro-social manner. Given the importance of social touch, we hypothesized that active stroking elicits more sensory pleasure when touching others' skin than when touching one's own skin. In a set of six experiments (total N = 133) we found that healthy participants, mostly tested in pairs to account for any objective differences in skin softness, consistently judged another's skin as feeling softer and smoother than their own skin. We further found that this softness illusion appeared selectively when the touch activated a neurophysiological system for affective touch in the receiver. We conclude that this sensory illusion underlies a novel, bodily mechanism of socio-affective bonding and enhances our motivation to touch others.

Doing good or bad: How interactions between action and emotion expectations shape the sense of agency.

The emotional consequences of our own and others' actions can influence our agentive self-awareness in social contexts. Positive outcomes are usually linked to the self and used for self-enhancement, whereas negative outcomes are more often attributed to others. In most situations, these causal attribution tendencies seem to be immediately present instead of involving reflective interpretations of the action experience. To address the question at which level of the cognitive hierarchy emotions and action perception interact, we adopted a social reward anticipation paradigm. Here, participants or their interaction partner received positive or negative action outcomes and performed speeded attribution choices regarding causation of the action outcome. Event-Related Potential (ERP) results showed that the emotional value of an outcome already influenced the classical N1 self-attenuation effect, with reduced embodied agentive self-awareness for negative outcomes at initial sensorimotor stages. At the level of the N300, the degree of updating and affective evaluation associated with the respective attributive decision was reflected and particularly associated to attribution tendencies for positive events. Our results show an early interaction between emotion and agency processes, and suggest that self-serving cognition can be grounded in embodied knowledge from low-level sensorimotor mechanisms.

Affective coding: the emotional dimension of agency.

The sense of agency (SoA) (i.e., the registration that I am the initiator and controller of my actions and relevant events) is associated with several affective dimensions. This makes it surprising that the emotion factor has been largely neglected in the field of agency research. Current empirical investigations of the SoA mainly focus on sensorimotor signals (i.e., efference copy) and cognitive cues (i.e., intentions, beliefs) and on how they are integrated. Here we argue that this picture is not sufficient to explain agency experience, since agency and emotions constantly interact in our daily life by several ways. Reviewing first recent empirical evidence, we show that self-action perception is in fact modulated by the affective valence of outcomes already at the sensorimotor level. We hypothesize that the "affective coding" between agency and action outcomes plays an essential role in agency processing, i.e., the prospective, immediate or retrospective shaping of agency representations by affective components. This affective coding of agency be differentially altered in various neuropsychiatric diseases (e.g., schizophrenia vs. depression), thus helping to explain the dysfunctions and content of agency experiences in these diseases.

Dysfunctional forward model mechanisms and aberrant sense of agency in obsessive-compulsive disorder.

BACKGROUND: Patients with obsessive-compulsive disorder (OCD) often lack the experience of action completion and have altered feelings of agency. This might depend on the integrity of predictions of action outcomes generated by forward models of the motor system. Such motor predictions are critical for inhibitory gating of actions and their consequences. Therefore, it was hypothesized that OCD patients show compromised forward model mechanisms. METHODS: To test whether inhibitory gating based on motor predictions is altered in OCD, we used electroencephalography to measure suppression of the N1 component of the event-related potential during active generation and passive observation of visual feedback in 18 OCD patients and 18 healthy control subjects. Predictability of action feedback was manipulated on the basis of action and external cues, and simultaneous agency judgments were assessed. RESULTS: OCD patients showed significantly reduced N1 suppression to actively generated feedback as compared with passively observed feedback. Moreover, in OCD patients, the N1 was not modulated by additional predictive motor cues as observed in control subjects. Patients also reported enhanced estimations of agency experience, which correlated with the strength of incompleteness feelings. CONCLUSIONS: OCD patients fail to predict and suppress the sensory consequences of their own actions. The increased mismatch between expected and actual outcomes caused by this forward model dysfunction may explain persistent feelings of incompleteness even after properly executed actions and the augmented search for control in these patients.