The blushing brain: neural substrates of cheek temperature increase in response to self-observation.

Darwin proposed that blushing-the reddening of the face owing to heightened self-awareness-is 'the most human of all expressions'. Yet, relatively little is known about the underlying mechanisms of blushing. Theories diverge on whether it is a rapid, spontaneous emotional response that does not involve reflection upon the self or whether it results from higher-order socio-cognitive processes. Investigating the neural substrates of blushing can shed light on the mental processes underlying blushing and the mechanisms involved in self-awareness. To reveal neural activity associated with blushing, 16-20 year-old participants (n = 40) watched pre-recorded videos of themselves (versus other people as a control condition) singing karaoke in a magnetic resonance imaging scanner. We measured participants' cheek temperature increase-an indicator of blushing-and their brain activity. The results showed that blushing is higher when watching oneself versus others sing. Those who blushed more while watching themselves sing had, on average, higher activation in the cerebellum (lobule V) and the left paracentral lobe and exhibited more time-locked processing of the videos in early visual cortices. These findings show that blushing is associated with the activation of brain areas involved in emotional arousal, suggesting that it may occur independently of higher-order socio-cognitive processes. Our results provide new avenues for future research on self-awareness in infants and non-human animals.

The state-trait sense of self inventory: A psychometric study of self-experience and its relation to psychosis-like manifestations.

The sense of self is a fundamental construct in the study of the mind, yet its psychological nature remains elusive. We introduce a novel 25-item inventory to investigate selfhood both as an enduring trait and a temporary state. We hypothesized two foundational aspects of the self: identity (related to self-referencing and continuity over time) and agency (the perception of controlling own's actions and thoughts). Results from two population studies highlight a singular self-trait factor combining agency and identity. In contrast, self-state measures revealed a bifactorial structure with a high-order factor and three lower-order subfactors: state-identity, state-agency, and state-technology. These factors were predictive of psychosis-like experiences, schizotypal traits, and hopelessness. Mediation analysis demonstrated that the negative association between the sense of self and hopelessness is mediated by depressive manifestations. Our research provides a tool to shed new light on the complexity of the sense of self and its mental health implications.

Brain and behavioral contributions to individual choices in response to affective-cognitive persuasion.

Affective and cognitive information conveyed by persuasive stimuli is evaluated and integrated by individuals according to their behavioral predispositions. However, the neurocognitive structure that supports persuasion based on either affective or cognitive content is poorly understood. Here, we examine the neural and behavioral processes supporting choices based on affective and cognitive persuasion by integrating 4 information processing features: intrinsic brain connectivity, stimulus-evoked brain activity, intrinsic affective-cognitive orientation, and explicit target evaluations. We found that the intrinsic cross-network connections of a multimodal fronto-parietal network are associated with individual affective-cognitive orientation. Moreover, using a cross-validated classifier, we found that individuals' intrinsic brain-behavioral dimensions, such as affective-cognitive orientation and intrinsic brain connectivity, can predict individual choices between affective and cognitive targets. Our findings show that affective- and cognitive-based choices rely on multiple sources, including behavioral orientation, stimulus evaluation, and intrinsic functional brain architecture.

An integrative perspective on the role of touch in the development of intersubjectivity.

Touch concerns a fundamental component of sociality. In this review, we examine the hypothesis that somatomotor development constitutes a crucial psychophysiological element in the ontogeny of intersubjectivity. An interdisciplinary perspective is provided on how the communication channel of touch contributes to the sense of self and extends to the social self. During gestation, the transformation of random movements into organized sequences of actions with sensory consequences parallels the development of the brain's functional architecture. Brain subsystems shaped by the coordinated activity of somatomotor circuits to support these first body-environment interactions are the first brain functional arrangements to develop. We propose that tactile self-referring behaviour during gestation constitutes a prototypic mode of interpersonal exchange that supports the subsequent development of intersubjective exchange. The reviewed research suggests that touch constitutes a pivotal bodily experience that in early stages builds and later filters self-other interactions. This view is corroborated by the fact that aberrant social-affective touch experiences appear fundamentally associated with attachment anomalies, interpersonal trauma, and personality disorders. Given the centrality of touch for the development of intersubjectivity and for psychopathological conditions in the social domain, dedicated research is urged to elucidate the role of touch in the evolution of subjective self-other coding.

Testing the Magnitude of Correlations Across Experimental Conditions.

Correlation coefficients are often compared to investigate data across multiple research fields, as they allow investigators to determine different degrees of correlation to independent variables. Even with adequate sample size, such differences may be minor but still scientifically relevant. To date, although much effort has gone into developing methods for estimating differences across correlation coefficients, adequate tools for variable sample sizes and correlational strengths have yet to be tested. The present study evaluated four different methods for detecting the difference between two correlations and tested the adequacy of each method using simulations with multiple data structures. The methods tested were Cohen's q, Fisher's method, linear mixed-effects models (LMEM), and an ad hoc developed procedure that integrates bootstrap and effect size estimation. Correlation strengths and sample size was varied across a wide range of simulations to test the power of the methods to reject the null hypothesis (i.e., the two correlations are equal). Results showed that Fisher's method and the LMEM failed to reject the null hypothesis even in the presence of relevant differences between correlations and that Cohen's method was not sensitive to the data structure. Bootstrap followed by effect size estimation resulted in a fair, unbiased compromise for estimating quantitative differences between statistical associations and producing outputs that could be easily compared across studies. This unbiased method is easily implementable in MatLab through the bootes function, which was made available online by the author at MathWorks.

Appropriately Tuning Stochastic-Psychometric Properties of the Balloon Analog Risk Task.

The Balloon Analog Risk Task (BART) allows to experimentally assess individuals' risk-taking profiles in an ecologically sound setting. Many psychological and neuroscientific studies implemented the BART for its simplicity and intuitive nature. However, some issues in the design of the BART are systematically unconsidered in experimental paradigms, which may bias the estimation of individual risk-taking profiles. Since there are no methodological guidelines for implementing the BART, many variables (e.g., the maximum explosion probabilities, the rationale underlying stochastic events) vary inconstantly across experiments, possibly producing contrasting results. Moreover, the standard version of the BART is affected by the interaction of an individual-dependent, unavoidable source of stochasticity with a trial-dependent, more ambiguous source of stochasticity (i.e., the probability of the balloon to explode). This paper shows the most appropriate experimental choices for having the lowest error in the approximation of risk-taking profiles. Performance tests of a series of simulated data suggest that a more controlled, eventually non-stochastic version of the BART, better approximates original risk-taking profiles. Selecting optimal BART parameters is particularly important in neuroscience experiments to optimize the number of trials in a time window appropriate for acquiring neuroimaging data. We also provide helpful suggestions to researchers in many fields to allow the implementation of optimized risk-taking experiments using the BART.

Probabilistically Weighted Multilayer Networks disclose the link between default mode network instability and psychosis-like experiences in healthy adults.

The brain is a complex system in which the functional interactions among its subunits vary over time. The trajectories of this dynamic variation contribute to inter-individual behavioral differences and psychopathologic phenotypes. Despite many methodological advancements, the study of dynamic brain networks still relies on biased assumptions in the temporal domain. The current paper has two goals. First, we present a novel method to study multilayer networks: by modelling intra-nodal connections in a probabilistic, biologically driven way, we introduce a temporal resolution of the multilayer network based on signal similarity across time series. This new method is tested on synthetic networks by varying the number of modules and the sources of noise in the simulation. Secondly, we implement these probabilistically weighted (PW) multilayer networks to study the association between network dynamics and subclinical, psychosis-relevant personality traits in healthy adults. We show that the PW method for multilayer networks outperforms the standard procedure in modular detection and is less affected by increasing noise levels. Additionally, the PW method highlighted associations between the temporal instability of default mode network connections and psychosis-like experiences in healthy adults. PW multilayer networks allow an unbiased study of dynamic brain functioning and its behavioral correlates.

Intrinsic Shapes of Empathy: Functional Brain Network Topology Encodes Intersubjective Experience and Awareness Traits.

Trait empathy is an essential personality feature in the intricacy of typical social inclinations of individuals. Empathy is likely supported by multilevel neuronal network functioning, whereas local topological properties determine network integrity. In the present functional MRI study (N = 116), we aimed to trace empathic traits to the intrinsic brain network architecture. Empathy was conceived as composed of two dimensions within the concept of pre-reflective, intersubjective understanding. Vicarious experience consists of the tendency to resonate with the feelings of other individuals, whereas intuitive understanding refers to a natural awareness of others' emotional states. Analyses of graph theoretical measures of centrality showed a relationship between the fronto-parietal network and psychometric measures of vicarious experience, whereas intuitive understanding was associated with sensorimotor and subcortical networks. Salience network regions could constitute hubs for information processing underlying both dimensions. The network properties related to empathy dimensions mainly concern inter-network information flow. Moreover, interaction effects implied several sex differences in the relationship between functional network organization and trait empathy. These results reveal that distinct intrinsic topological network features explain individual differences in separate dimensions of intersubjective understanding. The findings could help understand the impact of brain damage or stimulation through alterations of empathy-related network integrity.

Combining local and global evolutionary trajectories of brain-behaviour relationships through game theory.

The study of the evolution of brain-behaviour relationships concerns understanding the causes and repercussions of cross- and within-species variability. Understanding such variability is a main objective of evolutionary and cognitive neuroscience, and it may help explaining the appearance of psychopathological phenotypes. Although brain evolution is related to the progressive action of selection and adaptation through multiple paths (e.g. mosaic vs. concerted evolution, metabolic vs. structural and functional constraints), a coherent, integrative framework is needed to combine evolutionary paths and neuroscientific evidence. Here, we review the literature on evolutionary pressures focusing on structural-functional changes and developmental constraints. Taking advantage of recent progress in neuroimaging and cognitive neuroscience, we propose a twofold hypothetical model of brain evolution. Within this model, global and local trajectories imply rearrangements of neural subunits and subsystems and of behavioural repertoires of a species, respectively. We incorporate these two processes in a game in which the global trajectory shapes the structural-functional neural substrates (i.e. players), while the local trajectory shapes the behavioural repertoires (i.e. stochastic payoffs).

The Prospective Sense of Agency is Rooted in Local and Global Properties of Intrinsic Functional Brain Networks.

The sense of agency (SoA) refers to a constitutional aspect of the self describing the extent to which individuals feel in control over their actions and consequences thereof. Although the SoA has been associated with mental health and well-being, it is still unknown how interindividual variability in the SoA is embedded in the intrinsic brain organization. We hypothesized that the prospective component of an implicit SoA is associated with brain networks related to SoA and sensorimotor predictions on multiple spatial scales. We replicated previous findings by showing a significant prospective SoA as indicated by intentional binding effects. Then, using task-free fMRI and graph analysis, we analyzed associations between intentional binding effects and the intrinsic brain organization at regional, modular, and whole-brain scales. The results showed that intermodular connections of a frontoparietal module including the premotor cortex, supramarginal gyrus, and dorsal precuneus are associated with individual differences in prospective intentional binding. Notably, prospective intentional binding effects were also related to global brain modularity within a specific structural resolution range. These findings suggest that an implicit SoA generated through sensorimotor predictions relies on the intrinsic organization of the brain connectome on both local and global scales.

The evolving sense of agency: Context recency and quality modulate the interaction between prospective and retrospective processes.

Humans acquire a sense of agency through their interactions with the world and their sensory consequences. Previous studies have highlighted stable agency-related phenomena like intentional binding, which depend on both prospective, context-dependent and retrospective, outcome-dependent processes. In the current study, we investigated the interaction between prospective and retrospective processes underlying the adaptation of an ongoing sense of agency. The results showed that prospective intentional binding developed during a temporal window of up to 20 prior events was independent of the nature of the ongoing event. By contrast, the characteristics of the ongoing event retrospectively influenced prospective intentional binding developed during a temporal window narrower than 6 prior events. These findings characterize the interaction between prospective and retrospective mechanisms as a fundamental process to continuously update the sense of agency through sensorimotor learning. High psychosis-like experience traits weakened this interaction, suggesting that reduced adaption to the context contributes to altered self-experience.

I am Me: Brain systems integrate and segregate to establish a multidimensional sense of self.

Humans experience a sense of self, which is proposed to emerge from the integration of intrinsic and extrinsic self-processing through the propagation of information across brain systems. Using a novel functional magnetic resonance imaging (fMRI) paradigm, we tested this hypothesis in a non-clinical sample by modulating the intrinsic and extrinsic self-relatedness of auditory action consequences in terms of identity and agency, respectively. In addition, the relevance of individual traits associated with altered self-experiences (e.g., psychosis-like experiences) was examined. The task-evoked fMRI results showed distinctive associations between the neural coding of identity and negative affect traits, and between agency and psychosis-like experiences. Most importantly, regarding the functional connectivity analysis, graph theoretical measures demonstrated that the simultaneous processing of identity and agency relies on the functional integration and segregation of default mode, sensorimotor, language, and executive brain networks. Finally, cross-network interactions mediated by executive and sensorimotor regions were negatively associated with psychosis-like experiences when the intrinsic and extrinsic self-relatedness of action consequences conflicted. These findings provide evidence that the self is a multidimensional phenomenon rooted in the functional interactions between large-scale neuronal networks. Such interactions may have particular relevance for self-experience alterations.

Environmental control and psychosis-relevant traits modulate the prospective sense of agency in non-clinical individuals.

The sense of agency concerns the experience of being the source of one's own actions and their consequences. An altered sense of agency can occur due to task automation and in psychosis. We tested in a non-clinical sample the hypothesis that reducing voluntary task control diminishes intentional binding as an implicit indicator of the sense of agency, possibly interacting with psychosis-relevant personality traits. Agent-device interactions were manipulated obtaining positive-control (voluntary interaction), no-control (automation), and negative-control (device-commanded interaction) groups. The main results showed reduced prospective intentional binding (predictive coding of action consequences) in the no-control and negative-control groups, compared to the positive-control group. Psychosis-like experiences covaried positively with intentional binding in the no-control group, but negatively in the negative-control group. Moreover, positive-social traits were associated with increased intentional binding in the positive-control group. These findings demonstrate the interplay between environmental and individual differences variables in establishing the implicit sense of agency.

Spontaneous Brain Activity Predicts Task-Evoked Activity During Animate Versus Inanimate Touch.

The spontaneous activity of the brain is characterized by an elaborate temporal structure with scale-free properties as indexed by the power law exponent (PLE). We test the hypothesis that spontaneous brain activity modulates task-evoked activity during interactions with animate versus inanimate stimuli. For this purpose, we developed a paradigm requiring participants to actively touch either animate (real hand) or inanimate (mannequin hand) stimuli. Behaviorally, participants perceived the animate target as closer in space, temporally more synchronous with their own self, and more personally relevant, compared with the inanimate. Neuronally, we observed a modulation of task-evoked activity by animate versus inanimate interactions in posterior insula, in medial prefrontal cortex, comprising anterior cingulate cortex, and in medial superior frontal gyrus. Among these regions, an increased functional connectivity was shown between posterior insula and perigenual anterior cingulate cortex (PACC) during animate compared with inanimate interactions and during resting state. Importantly, PLE during spontaneous brain activity in PACC correlated positively with PACC task-evoked activity during animate versus inanimate stimuli. In conclusion, we demonstrate that brain spontaneous activity in PACC can be related to the distinction between animate and inanimate stimuli and thus might be specifically tuned to align our brain with its animate environment.

Brain network profiling defines functionally specialized cortical networks.

Neuroimaging research made rapid advances in the study of the functional architecture of the brain during the past decade. Many proposals endorsed the relevance of large-scale brain networks, defined as ensembles of brain regions that exhibit highly correlated signal fluctuations. However, analysis methods need further elaboration to define the functional and anatomical extent of specialized subsystems within classical networks with a high reliability. We present a novel approach to characterize and examine the functional proprieties of brain networks. This approach, labeled as brain network profiling (BNP), considers similarities in task-evoked activity and resting-state functional connectivity across biologically relevant brain subregions. To combine task-driven activity and functional connectivity features, principal components were extracted separately for task-related beta values and resting-state functional connectivity z-values (data available from the Human Connectome Project), from 360 brain parcels. Multiple clustering procedures were employed to assess if different clustering methods (Gaussian mixtures; k-means) and/or data structures (task and rest data; only rest data) led to improvements in the replication of the brain architecture. The results indicated that combining information from resting-state functional connectivity and task-evoked activity and using Gaussian mixtures models for clustering produces more reliable results (99% replication across data sets). Moreover, the findings revealed a high-resolution partition of the cerebral cortex in 16 networks with unique functional connectivity and/or task-evoked activity profiles. BNP potentially offers new approaches to advance the investigation of the brain functional architecture.

Functional connections between activated and deactivated brain regions mediate emotional interference during externally directed cognition.

Recent evidence shows that task-deactivations are functionally relevant for cognitive performance. Indeed, higher cognitive engagement has been associated with higher suppression of activity in task-deactivated brain regions - usually ascribed to the Default Mode Network (DMN). Moreover, a negative correlation between these regions and areas actively engaged by the task is associated with better performance. DMN regions show positive modulation during autobiographical, social, and emotional tasks. However, it is not clear how processing of emotional stimuli affects the interplay between the DMN and executive brain regions. We studied this interplay in an fMRI experiment using emotional negative stimuli as distractors. Activity modulations induced by the emotional interference of negative stimuli were found in frontal, parietal, and visual areas, and were associated with modulations of functional connectivity between these task-activated areas and DMN regions. A worse performance was predicted both by lower activity in the superior parietal cortex and higher connectivity between visual areas and frontal DMN regions. Connectivity between right inferior frontal gyrus and several DMN regions in the left hemisphere was related to the behavioral performance. This relation was weaker in the negative than in the neutral condition, likely suggesting less functional inhibitions of DMN regions during emotional processing. These results show that both executive and DMN regions are crucial for the emotional interference process and suggest that DMN connections are related to the interplay between externally-directed and internally-focused processes. Among DMN regions, superior frontal gyrus may be a key node in regulating the interference triggered by emotional stimuli.