Reputation risk during dishonest social decision-making modulates anterior insular and cingulate cortex activity and connectivity.

To explore the neural underpinnings of (dis)honest decision making under quasi-ecological conditions, we used an fMRI adapted version of a card game in which deceptive or truthful decisions are made to an opponent, with or without the risk of getting caught by them. Dishonest decisions were associated to increased activity in a cortico-subcortical circuit including the bilateral anterior cingulate cortex (ACC), anterior insula (AI), left dorsolateral prefrontal cortex, supplementary motor area, and right caudate. Crucially, deceptive immoral decisions under reputation risk enhanced activity of - and functional connectivity between - the bilateral ACC and left AI, suggesting the need for heightened emotional processing and cognitive control when making immoral decisions under reputation risk. Tellingly, more manipulative individuals required less involvement of the ACC during risky self-gain lies but more involvement during other-gain truths, pointing to the need of cognitive control only when going against one's own moral code.

Re-cognizing the new self: The neurocognitive plasticity of self-processing following facial transplantation.

The face is a defining feature of our individuality, crucial for our social interactions. But what happens when the face connected to the self is radically altered or replaced? We address the plasticity of self-face recognition in the context of facial transplantation. While the acquisition of a new face following facial transplantation is a medical fact, the experience of a new identity is an unexplored psychological outcome. We traced the changes in self-face recognition before and after facial transplantation to understand if and how the transplanted face gradually comes to be perceived and recognized as the recipient's own new face. Neurobehavioral evidence documents a strong representation of the pre-injury appearance pre-operatively, while following the transplantation, the recipient incorporates the new face into his self-identity. The acquisition of this new facial identity is supported by neural activity in medial frontal regions that are considered to integrate psychological and perceptual aspects of the self.

From the Viscera to First Impressions: Phase-Dependent Cardio-Visual Signals Bias the Perceived Trustworthiness of Faces.

When we see new people, we rapidly form first impressions. Whereas past research has focused on the role of morphological or emotional cues, we asked whether transient visceral states bias the impressions we form. Across three studies (N = 94 university students), we investigated how fluctuations of bodily states, driven by the interoceptive impact of cardiac signals, influence the perceived trustworthiness of faces. Participants less often chose faces presented in synchrony with their own cardiac systole as more trustworthy than faces presented out of synchrony. Participants also explicitly judged faces presented in synchrony with their cardiac systole as less trustworthy. Finally, the presentation of faces in synchrony with participants' cardiac diastole did not modulate participants' perceptions of the faces' trustworthiness, suggesting that the systolic phase is necessary for such interoceptive effects. These findings highlight the role of phasic interoceptive information in the processing of social information and provide a mechanistic account of the role of visceroception for social perception.

Event-related transcutaneous vagus nerve stimulation modulates behaviour and pupillary responses during an auditory oddball task.

Transcutaneous auricular vagus nerve stimulation (taVNS) is a neuromodulatory technique that is thought to activate the Locus Coeruleus-Noradrenaline (LC-NA) system. Standard taVNS protocols consist of the administration of intermittent or continuous stimulation over long periods. However, there is currently a limited understanding of the temporal dynamics of taVNS modulation of cognitive processes, as well as its mechanisms of action. We argue that novel stimulation approaches, informed by established theories of the LC-NA system, are needed to further our understanding of the neurocognitive underpinnings of taVNS. In this pre-registered study, we tested whether an "event-related" taVNS protocol can modulate the LC-NA system. In a within-subject design (single session) we delivered brief trains of taVNS (3 s) during an auditory oddball paradigm. The taVNS was time-locked to the target stimuli presentation and randomly interleaved with sham stimulation. Response times (RT) and stimuli-driven pupillary diameter (PD) were used as indices of LC-NA activity. Results revealed that active taVNS increased RT to targets, as compared to sham trials. Notably, in line with current theories of LC-NA functioning, taVNS modulation of target-related pupil dilation depended on pre-stimulation PD, an index of baseline LC-NA activity. In particular, active (vs. sham) taVNS was associated with smaller pupil dilation in trials where the baseline PD was small. These results demonstrate, for the first time, the effectiveness of brief event-related taVNS in the modulation of cognitive processes and highlight the importance of using pupil size as an index of tonic and phasic LC-NA activity.

Neural divergence and convergence for attention to and detection of interoceptive and somatosensory stimuli.

Body awareness is constructed by signals originating from within and outside the body. How do these apparently divergent signals converge? We developed a signal detection task to study the neural convergence and divergence of interoceptive and somatosensory signals. Participants focused on either cardiac or tactile events and reported their presence or absence. Beyond some evidence of divergence, we observed a robust overlap in the pattern of activation evoked across both conditions in frontal areas including the insular cortex, as well as parietal and occipital areas, and for both attention and detection of these signals. Psycho-physiological interaction analysis revealed that right insular cortex connectivity was modulated by the conscious detection of cardiac compared to somatosensory sensations, with greater connectivity to occipito-parietal regions when attending to cardiac signals. Our findings speak in favour of the inherent convergence of bodily-related signals and move beyond the apparent antagonism between exteroception and interoception.

Emotion matters: Different psychophysiological responses to expressive and non-expressive full-body movements.

We explore dance video clip stimuli as a means to test human observers' accuracy in detecting genuine emotional expressivity in full-body movements. Stimuli of every-day-type full-body expressions of emotions usually use culturally very recognizable actions (e.g. fist shaking for anger, etc). However, expressive dance movement stimuli can be created to contain fully abstract movements. The expressivity results from subtle variations in the body movements of the expressor, and emotions cannot be recognised by observers via particular actions (e.g. fist shaking, etc). Forty-one participants watched and rated 24 pairs of short dance videos -from a published normalised dance stimuli library- in randomised order (N = 48). Of each carefully matched pair, one version of the full-body movement sequence had been danced to be emotionally genuinely expressive (clip a), while the other version of the same sequence (clip b) had been danced -while technically correct- without any emotional expressivity. Participants rated (i) expressivity (to test their accuracy; block 1), and (ii) how much they liked each movement (an implicit measure to test their emotional response ("liking"); block 2). Participants rated clips that were intended to be expressive as more expressive (part 1: expressivity ratings), and liked those expressive clips more than the non-expressive clips (part 2: liking ratings). Besides, their galvanic skin response differed, depending on the category of clips they were watching (expressive vs. non-expressive), and this relationship was modulated by interceptive accuracy and arts experience. Results are discussed in relation to the Body Precision Hypothesis and the Hypothesis of Constructed Emotion.

Interoceptive Awareness Is Negatively Related to the Exteroceptive Manipulation of Bodily Self-Location.

The perception of being located within one's body (i.e., bodily self-location) is an essential feature of everyday self-experience. However, by manipulating exteroceptive input, healthy participants can easily be induced to perceive themselves as being spatially dislocated from their physical bodies. It has previously been suggested that interoception, i.e., the processing of inner physiological signals, contributes to the stability of body representations; however, this relationship has not previously been tested for different dimensions of interoception and bodily self-location. In the present study, using an advanced automatized setup, we systematically manipulated participants' perspective of their own body (first- vs third-person perspective) as well as the synchrony of visuotactile stimulation (synchronous vs asynchronous). The malleability of bodily self-location was assessed using a questionnaire targeting in-body and out-of-body experiences. Participants also performed a heartbeat discrimination task to assess their interoceptive accuracy (behavioral performance), interoceptive sensibility (confidence in their interoceptive abilities), and interoceptive awareness (meta-cognitive representation of interoceptive signals). Bodily self-location was significantly influenced by perspective, with third-person perspective being associated with stronger out-of-body experiences compared to first-person perspective. Furthermore, there was a significant perspective × stimulation interaction, with subsequent analyses showing that participants reported out-of-body experiences particularly under third-person perspective combined with synchronous visuotactile stimulation. Correlation and regression analyses revealed that meta-cognitive interoceptive awareness was specifically and negatively related to the exteroceptively mediated malleability of body experiences. These results indicate that the perception of the self being located within one's body relies on the interaction of exteroceptive input and higher-order interoceptive abilities. This has implications for theoretical considerations about the bodily self in health as well as for the understanding of disturbed bodily self-processing in clinical contexts.

Seeing myself through my heart: Cortical processing of a single heartbeat speeds up self-face recognition.

Recent research has highlighted the contribution of interoceptive signals to different aspects of bodily self-consciousness (BSC) by means of the cardio-visual stimulation - i.e. perceiving a pulsing stimulus in synchrony with one's own heart. Here, for the first time, we investigate the effects of individual heartbeats on a critical feature of BSC, namely the recognition of one's own face. Across two studies, we explored the cardiac-timing effects on a classic self-face recognition task. In Study 1, participants saw morphed faces that contained different percentages of the self-face and that of another unfamiliar individual. Study 2 used a similar design, albeit participants saw morphed faces of the self-face and that of a familiar other to provide a better control of self-familiarity. Results from both studies consistently revealed that the cortical processing of cardiac afferent signals conveyed by the firing of arterial baroreceptors affects the speed, but not the accuracy, of self-face recognition, when a single picture is presented during cardiac systole, as compared to diastole. This effect is stronger and more stable for stimuli with more self-cues than other-cues and for 'ambiguous' stimuli - i.e. at the individual point of subjective equality. Results from Study 2 also revealed that cardiac effects on the speed of self-face recognition cannot be explained simply on the basis of the imbalanced familiarity between the self's and other's faces used. The present findings highlight the interoceptive contributions to self-recognition and may be expand our understanding of pathological disturbances of self-experience.

Not on my team: Medial prefrontal cortex responses to ingroup fusion and unfair monetary divisions.

OBJECTIVE: People are highly attuned to fairness, with people willingly suffering personal costs to prevent others benefitting from unfair acts. Are fairness judgments influenced by group alignments? A new theory posits that we favor ingroups and denigrate members of rival outgroups when our personal identity is fused to a group. Although the mPFC has been separately implicated in group membership and fairness processing, it is unclear whether group alignments affect medial prefrontal cortex (mPFC) activity in response to fairness. Here, we examine the contribution of different regions of the mPFC to processing from ingroup and outgroup members and test whether its response differs depending on how fused we are to an ingroup. METHODS: Subjects performed rounds of the Ultimatum Game, being offered fair or unfair divisions of money from supporters of the same soccer team (ingroup), the fiercest rival (outgroup) or neutral individuals whilst undergoing functional Magnetic Resonance Imaging (fMRI). RESULTS: Strikingly, people willingly suffered personal costs to prevent outgroup members benefitting from both unfair and fair offers. Activity across dorsal and ventral (VMPFC) portions of the mPFC reflected an interaction between fairness and group membership. VMPFC activity in particular was consistent with it coding one's fusion to a group, with the fairness by group membership interaction correlating with the extent that the responder's identity was fused to the ingroup. CONCLUSIONS: The influence of fusion on social behavior therefore seems to be linked to processing in the VMPFC.

Afferent cardiac signals modulate attentional engagement to low spatial frequency fearful faces.

Despite the growing consensus that the continuous dynamic cortical representations of internal bodily states shape the subjective experience of emotions, physiological arousal is typically considered only a consequence and rarely a determinant of the emotional experience. Recent experimental approaches study how afferent autonomic signals from the heart modulate the processing of sensory information by focussing on the phasic properties of arterial baroreceptor firing that is active during cardiac systole and quiescent during cardiac diastole. For example, baroreceptor activation has been shown to enhance the processing of threat-signalling stimuli. Here, we investigate the role of cardiac afferent signals in the rapid engagement and disengagement of attention to fear stimuli. In an adapted version of the emotional attentional cueing paradigm, we timed the presentation of cues, either fearful or neutral faces, to coincide with the different phases of the cardiac cycle. Moreover, we presented cues with different spatial frequency ranges to investigate how these interoceptive signals influence the processing of visual information. Results revealed a selective enhancement of attentional engagement to low spatial frequency fearful faces presented during cardiac systole relative to diastole. No cardiac cycle effects were observed to high spatial frequency nor broad spatial frequency cues. These findings expand our mechanistic understanding of how body-brain interactions may impact the visual processing of fearful stimuli and contribute to the increased attentional capture of threat signals.

Perceived warmth and competence of others shape voluntary deceptive behaviour in a morally relevant setting.

The temptation to deceive others compares to a moral dilemma: it involves a conflict between the temptation to obtain some benefit and the desire to conform to personal and social moral norms or avoid aversive social consequences. Thus, people might feel different levels of emotional and moral conflict depending on the target of the deception. Here we explored, in a morally relevant setting, how social judgements based on two fundamental dimensions of human social cognition - 'warmth' and 'competence' - impact on the decision to deceive others. Results revealed independent effects for warmth and competence. Specifically, while people are less inclined to deceive for self-gain those individuals they perceive as warm, they also tend to lie more to highly competent others. Furthermore, the perceived warmth and competence modulated the general tendency to reduce deceptive behaviour when there was a risk of disclosure compared to when the lying was anonymous, highlighting the importance of these judgements in social evaluation processes. Together, our results demonstrate that the emotional costs and personal moral standards that inhibit engagement in deceptive behaviour are not stable but rather malleable according to the target and the consequences of the deception.

Heartfelt Self: Cardio-Visual Integration Affects Self-Face Recognition and Interoceptive Cortical Processing.

The sense of body-ownership relies on the representation of both interoceptive and exteroceptive signals coming from one's body. However, it remains unknown how the integration of bodily signals coming from "outside" and "inside" the body is instantiated in the brain. Here, we used a modified version of the Enfacement Illusion to investigate whether the integration of visual and cardiac information can alter self-face recognition (Experiment 1) and neural responses to heartbeats (Experiment 2). We projected a pulsing shade, that was synchronous or asynchronous with the participant's heartbeat, onto a picture depicting the participant's face morphed with the face of an unfamiliar other. Results revealed that synchronous (vs. asynchronous) cardio-visual stimulation led to increased self-identification with the other's face (Experiment 1), while during stimulation, synchronicity modulated the amplitude of the Heartbeat Evoked Potential, an electrophysiological index of cortical interoceptive processing (Experiment 2). Importantly, the magnitude of the illusion-related effects was dependent on, and increased linearly, with the participants' Interoceptive Accuracy. These results provide the first direct neural evidence for the integration of interoceptive and exteroceptive signals in bodily self-awareness.

Cardiac afferent activity modulates the expression of racial stereotypes.

Negative racial stereotypes tend to associate Black people with threat. This often leads to the misidentification of harmless objects as weapons held by a Black individual. Yet, little is known about how bodily states impact the expression of racial stereotyping. By tapping into the phasic activation of arterial baroreceptors, known to be associated with changes in the neural processing of fearful stimuli, we show activation of race-threat stereotypes synchronized with the cardiovascular cycle. Across two established tasks, stimuli depicting Black or White individuals were presented to coincide with either the cardiac systole or diastole. Results show increased race-driven misidentification of weapons during systole, when baroreceptor afferent firing is maximal, relative to diastole. Importantly, a third study examining the positive Black-athletic stereotypical association fails to demonstrate similar modulations by cardiac cycle. We identify a body-brain interaction wherein interoceptive cues can modulate threat appraisal and racially biased behaviour in context-dependent ways.

Art reception as an interoceptive embodied predictive experience.

In the Distancing-Embracing model, an explanation is proposed for the apparent paradox that is the enjoyment of negative emotional states in art reception. Here, we argue for the advantages of grounding the psychological dynamics described in the model in established and empirically testable frameworks of brain functioning by thinking of art reception as an embodied experience guided by predictive coding.

Participants' above-chance recognition of own-heart sound combined with poor metacognitive awareness suggests implicit knowledge of own heart cardiodynamics.

Mounting evidence suggests that interoceptive signals are fundamentally important for the experience of the self. Thus far, studies on interoception have mainly focused on the ability to monitor the timing of ongoing heartbeats and on how these influence emotional and self-related processes. However, cardiac afferent signalling is not confined to heartbeat timing and several other cardiac parameters characterize cardiodynamic functioning. Building on the fact that each heart has its own self-specific cardio-dynamics, which cannot be expressed uniquely by heart rate, we devised a novel task to test whether people could recognize the sound of their own heart even when perceived offline and thus not in synchrony with ongoing heartbeats. In a forced-choice paradigm, participants discriminated between sounds of their own heartbeat (previously recorded with a Doppler device) versus another person's heart. Participants identified the sound of their own heart above chance, whereas their metacognition of performance - as calculated by contrasting performance against ratings of confidence - was considerably poorer. These results suggest an implicit access to fine-grained neural representations of elementary cardio-dynamic parameters beyond heartbeat timing.

Cardio-visual integration modulates the subjective perception of affectively neutral stimuli.

Interoception, which refers to the perception of internal body signals, has been consistently associated with emotional processing and with the sense of self. However, its influence on the subjective appraisal of affectively neutral and body-unrelated stimuli is still largely unknown. Across two experiments we sought to investigate this issue by asking participants to detect changes in the flashing rhythm of a simple stimulus (a circle) that could either be pulsing synchronously with their own heartbeats or following the pattern of another person's heart. While overall task performance did not vary as a function of cardio-visual synchrony, participants were better at identifying trials in which no change occurred when the flashes were synchronous with their own heartbeats. This study adds to the growing body of research indicating that we use our body as a reference point when perceiving the world; and extends this view by focusing on the role that signals coming from inside the body, such as heartbeats, may play in this referencing process. Specifically we show that private interoceptive sensations can be combined with affectively neutral information unrelated to the self to influence the processing of a multisensory percept. Results are discussed in terms of both standard multisensory integration processes and predictive coding theories.

Listening to your heart and feeling yourself: effects of exposure to interoceptive signals during the ultimatum game.

The ultimatum game (UG) is commonly used to study the tension between financial self-interest and social equity motives. Here, we investigated whether experimental exposure to interoceptive signals influences participants' behavior in the UG. Participants were presented with various bodily sounds--i.e., their own heart, another person's heart, or the sound of footsteps--while acting both in the role of responder and proposer. We found that listening to one's own heart sound, compared to the other bodily sounds: (1) increased subjective feelings of unfairness, but not rejection behavior, in response to unfair offers and (2) increased the unfair offers while playing in the proposer role. These findings suggest that heightened feedback of one's own visceral processes may increase a self-centered perspective and drive socioeconomic exchanges accordingly. In addition, this study introduces a valuable procedure to manipulate online the access to interoceptive signals and for exploring the interplay between viscero-sensory information and cognition.

Their pain is not our pain: brain and autonomic correlates of empathic resonance with the pain of same and different race individuals.

Recent advances in social neuroscience research have unveiled the neurophysiological correlates of race and intergroup processing. However, little is known about the neural mechanisms underlying intergroup empathy. Combining event-related fMRI with measurements of pupil dilation as an index of autonomic reactivity, we explored how race and group membership affect empathy-related responses. White and Black subjects were presented with video clips depicting white, black, and unfamiliar violet-skinned hands being either painfully penetrated by a syringe or being touched by a Q-tip. Both hemodynamic activity within areas known to be involved in the processing of first and third-person emotional experiences of pain, i.e., bilateral anterior insula, and autonomic reactivity were greater for the pain experienced by own-race compared to that of other-race and violet models. Interestingly, greater implicit racial bias predicted increased activity within the left anterior insula during the observation of own-race pain relative to other-race pain. Our findings highlight the close link between group-based segregation and empathic processing. Moreover, they demonstrate the relative influence of culturally acquired implicit attitudes and perceived similarity/familiarity with the target in shaping emotional responses to others' physical pain.

Epistasis between 5-HTTLPR and ADRA2B polymorphisms influences attentional bias for emotional information in healthy volunteers.

Individual differences in emotional processing are likely to contribute to vulnerability and resilience to emotional disorders such as depression and anxiety. Genetic variation is known to contribute to these differences but they remain incompletely understood. The serotonin transporter (5-HTTLPR) and α2B-adrenergic autoreceptor (ADRA2B) insertion/deletion polymorphisms impact on two separate but interacting monaminergic signalling mechanisms that have been implicated in both emotional processing and emotional disorders. Recent studies suggest that the 5-HTTLPR s allele is associated with a negative attentional bias and an increased risk of emotional disorders. However, such complex behavioural traits are likely to exhibit polygenicity, including epistasis. This study examined the contribution of the 5-HTTLPR and ADRA2B insertion/deletion polymorphisms to attentional biases for aversive information in 94 healthy male volunteers and found evidence of a significant epistatic effect (p<0.001). Specifically, in the presence of the 5-HTTLPR s allele, the attentional bias for aversive information was attenuated by possession of the ADRA2B deletion variant whereas in the absence of the s allele, the bias was enhanced. These data identify a cognitive mechanism linking genotype-dependent serotonergic and noradrenergic signalling that is likely to have implications for the development of cognitive markers for depression/anxiety as well as therapeutic drug effects and personalized approaches to treatment.

Deletion variant of alpha2b-adrenergic receptor gene moderates the effect of COMT val(158)met polymorphism on episodic memory performance.

The COMT val(158) variant has been associated with impaired cognitive function compared to the met(158) variant yet gene-gene interactions are not well described. In this study we demonstrate an interaction between this COMT polymorphism and a deletion variant of ADRA2B, the gene encoding the alpha2b-adrenergic receptor on episodic memory performance. Specifically, carriage of the ADRA2B deletion abolished the relative memory impairment in homozygous COMT val(158) carriers compared to met(158) carriers.