Dissecting abstract, modality-specific and experience-dependent coding of affect in the human brain.

Emotion and perception are tightly intertwined, as affective experiences often arise from the appraisal of sensory information. Nonetheless, whether the brain encodes emotional instances using a sensory-specific code or in a more abstract manner is unclear. Here, we answer this question by measuring the association between emotion ratings collected during a unisensory or multisensory presentation of a full-length movie and brain activity recorded in typically developed, congenitally blind and congenitally deaf participants. Emotional instances are encoded in a vast network encompassing sensory, prefrontal, and temporal cortices. Within this network, the ventromedial prefrontal cortex stores a categorical representation of emotion independent of modality and previous sensory experience, and the posterior superior temporal cortex maps the valence dimension using an abstract code. Sensory experience more than modality affects how the brain organizes emotional information outside supramodal regions, suggesting the existence of a scaffold for the representation of emotional states where sensory inputs during development shape its functioning.

Balancing the Senses: Electrophysiological Responses Reveal the Interplay between Somatosensory and Visual Processing During Body-Related Multisensory Conflict.

In the study of bodily awareness, the predictive coding theory has revealed that our brain continuously modulates sensory experiences to integrate them into a unitary body representation. Indeed, during multisensory illusions (e.g., the rubber hand illusion, RHI), the synchronous stroking of the participant's concealed hand and a fake visible one creates a visuotactile conflict, generating a prediction error. Within the predictive coding framework, through sensory processing modulation, prediction errors are solved, inducing participants to feel as if touches originated from the fake hand, thus ascribing the fake hand to their own body. Here, we aimed to address sensory processing modulation under multisensory conflict, by disentangling somatosensory and visual stimuli processing that are intrinsically associated during the illusion induction. To this aim, we designed two EEG experiments, in which somatosensory- (SEPs; Experiment 1; N = 18; F = 10) and visual-evoked potentials (VEPs; Experiment 2; N = 18; F = 9) were recorded in human males and females following the RHI. Our results show that, in both experiments, ERP amplitude is significantly modulated in the illusion as compared with both control and baseline conditions, with a modality-dependent diametrical pattern showing decreased SEP amplitude and increased VEP amplitude. Importantly, both somatosensory and visual modulations occur in long-latency time windows previously associated with tactile and visual awareness, thus explaining the illusion of perceiving touch at the sight location. In conclusion, we describe a diametrical modulation of somatosensory and visual processing as the neural mechanism that allows maintaining a stable body representation, by restoring visuotactile congruency under the occurrence of multisensory conflicts.

Modulation of Motor Awareness: A Transcranial Magnetic Stimulation Study in the Healthy Brain.

Previous studies on the mechanisms underlying willed actions reported that the premotor cortex may be involved in the construction of motor awareness. However, its exact role is still under investigation. Here, we investigated the role of the dorsal premotor cortex (PMd) in motor awareness by modulating its activity applying inhibitory rTMS to PMd, before a specific motor awareness task (under three conditions: without stimulation, after rTMS and after Sham stimulation). During the task, subjects had to trace straight lines to a given target, receiving visual feedback of the line trajectories on a computer screen. Crucially, in most trials, the trajectories on the screen were deviated, and to produce straight lines, subjects had to correct their movements towards the opposite direction. After each trial, participants were asked to judge whether the line seen on the computer screen corresponded to the line actually drawn. Results show that participants in the No Stimulation condition did not recognize the perturbation until 14 degrees of deviation. Importantly, active, but not Sham, rTMS significantly modulated motor awareness, decreasing the amplitude of the angle at which participants became aware of the trajectory correction. These results suggest that PMd plays a crucial role in action self-monitoring.

Abnormal sense of agency in eating disorders.

The feeling of controlling one's own actions and, through them, impacting the external environment (i.e. Sense of Agency-SoA) can be relevant in the eating disorders (EDs) symptomatology. Yet, it has been poorly investigated. This study aims to implicitly assess SoA exploiting the Sensory Attenuation paradigm in two groups of EDs patients (Anorexia Nervosa Restrictive and Anorexia Nervosa Binge-Purging or Bulimia Nervosa) compared to a control group. We find that controls perceive self-generated stimuli as less intense than other-generated ones showing the classic pattern of sensory attenuation. By contrast, EDs patients show the opposite pattern, with self-generated perceived as more intense than other-generated stimuli. This result indicates an alteration of the implicit component of the feeling of control in EDs patients, thus suggesting a potential implication of these results for the clinical practice and the treatment of EDs symptomatology.

Can static Rorschach stimuli perceived as in motion affect corticospinal excitability?

It has been proposed that seeing human movement or activity (M), while trying to say what the static Rorschach inkblot design look like, is accompanied by Mirror Neuron System (MNS)-like mirroring activity in the brain. The present study aimed to investigate whether the Rorschach cards eliciting M responses could affect the excitability of the motor cortex by recording motor evoked potentials (MEPs) elicited by single-pulse TMS over the primary motor cortex (M1). We hypothesized that Rorschach inkblot stimuli triggering the viewer's experience of human movement would increase corticospinal excitability. Twenty-one healthy volunteers (15 women) participated in the preliminary experiment, while another different sample of twenty-two healthy participants (11 women) ranging in age from 21 to 41 years was enrolled in the main experiment. Our results showed that the Rorschach cards known to be associated with a high number of M responses elicited human movement both as automatic internal sensations and as verbal production of responses involving human movement. However, contrary to our hypothesis, the reported internal feeling of human movement had no corresponding physiological counterpart, as the amplitude of MEPs did not increase. Possible and innovative explanations for the involvement of bottom-up and top-down processes were provided.

The role of early attachment experiences in modulating defensive peripersonal space.

Selecting appropriate defensive behaviours for threats approaching the space surrounding the body (peripersonal space, PPS) is crucial for survival. The extent of defensive PPS is measured by recording the hand-blink reflex (HBR), a subcortical defensive response. Higher-order cortical areas involved in PPS representation exert top-down modulation on brainstem circuits subserving HBR. However, it is not yet known whether pre-existing models of social relationships (internal working models, IWM) originating from early attachment experiences influence defensive responses. We hypothesized that organized IWM ensure adequate top-down regulation of brainstem activity mediating HBR, whereas disorganized IWM are associated with altered response patterns. To investigate attachment-dependent modulation on defensive responses, we used the Adult Attachment Interview to determine IWM and recorded HBR in two sessions (with or without the neurobehavioral attachment system activated). As expected, the HBR magnitude in individuals with organized IWM was modulated by the threat proximity to the face, regardless of the session. In contrast, for individuals with disorganized IWM, attachment system activation enhances HBR regardless of the threat position, suggesting that triggering emotional attachment experiences magnifies the threatening valence of external stimuli. Our results indicate that the attachment system exerts a strong modulation on defensive responses and the magnitude of PPS.

Spatial proximity to others induces plastic changes in the neural representation of the peripersonal space.

Peripersonal space (PPS) is a highly plastic "invisible bubble" surrounding the body whose boundaries are mapped through multisensory integration. Yet, it is unclear how the spatial proximity to others alters PPS boundaries. Across five experiments (N = 80), by recording behavioral and electrophysiological responses to visuo-tactile stimuli, we demonstrate that the proximity to others induces plastic changes in the neural PPS representation. The spatial proximity to someone else's hand shrinks the portion of space within which multisensory responses occur, thus reducing the PPS boundaries. This suggests that PPS representation, built from bodily and multisensory signals, plastically adapts to the presence of conspecifics to define the self-other boundaries, so that what is usually coded as "my space" is recoded as "your space". When the space is shared with conspecifics, it seems adaptive to move the other-space away from the self-space to discriminate whether external events pertain to the self-body or to other-bodies.

A modality-independent proto-organization of human multisensory areas.

The processing of multisensory information is based upon the capacity of brain regions, such as the superior temporal cortex, to combine information across modalities. However, it is still unclear whether the representation of coherent auditory and visual events requires any prior audiovisual experience to develop and function. Here we measured brain synchronization during the presentation of an audiovisual, audio-only or video-only version of the same narrative in distinct groups of sensory-deprived (congenitally blind and deaf) and typically developed individuals. Intersubject correlation analysis revealed that the superior temporal cortex was synchronized across auditory and visual conditions, even in sensory-deprived individuals who lack any audiovisual experience. This synchronization was primarily mediated by low-level perceptual features, and relied on a similar modality-independent topographical organization of slow temporal dynamics. The human superior temporal cortex is naturally endowed with a functional scaffolding to yield a common representation across multisensory events.

Moving without sensory feedback: online TMS over the dorsal premotor cortex impairs motor performance during ischemic nerve block.

The study investigates the role of dorsal premotor cortex (PMd) in generating predicted sensory consequences of movements, i.e. corollary discharges. In 2 different sessions, we disrupted PMd and parietal hand's multisensory integration site (control area) with transcranial magnetic stimulation (TMS) during a finger-sequence-tapping motor task. In this TMS sham-controlled design, the task was performed with normal sensory feedback and during upper-limb ischemic nerve block (INB), in a time-window where participants moved without somatosensation. Errors and movement timing (objective measures) and ratings about movement perception (subjective measures) were collected. We found that INB overall worsens objective and subjective measures, but crucially in the PMd session, the absence of somatosensation together with TMS disruption induced more errors, less synchronized movements, and increased subjective difficulty ratings as compared with the parietal control session (despite a carryover effect between real and sham stimulation to be addressed in future studies). Contrarily, after parietal area interference session, when sensory information is already missing due to INB, motor performance was not aggravated. Altogether these findings suggest that the loss of actual (through INB) and predicted (through PMd disruption) somatosensory feedback degraded motor performance and perception, highlighting the crucial role of PMd in generating corollary discharge.

Multisensory-driven facilitation within the peripersonal space is modulated by the expectations about stimulus location on the body.

Compelling evidence from human and non-human studies suggests that responses to multisensory events are fastened when stimuli occur within the space surrounding the bodily self (i.e., peripersonal space; PPS). However, some human studies did not find such effect. We propose that these dissonant voices might actually uncover a specific mechanism, modulating PPS boundaries according to sensory regularities. We exploited a visuo-tactile paradigm, wherein participants provided speeded responses to tactile stimuli and rated their perceived intensity while ignoring simultaneous visual stimuli, appearing near the stimulated hand (VTNear) or far from it (VTFar; near the non-stimulated hand). Tactile stimuli could be delivered only to one hand (unilateral task) or to both hands randomly (bilateral task). Results revealed that a space-dependent multisensory enhancement (i.e., faster responses and higher perceived intensity in VTNear than VTFar) was present when highly predictable tactile stimulation induced PPS to be circumscribed around the stimulated hand (unilateral task). Conversely, when stimulus location was unpredictable (bilateral task), participants showed a comparable multisensory enhancement in both bimodal conditions, suggesting a PPS widening to include both hands. We propose that the detection of environmental regularities actively shapes PPS boundaries, thus optimizing the detection and reaction to incoming sensory stimuli.

Bodily self-recognition in patients with pathological embodiment.

The ability to discriminate between one's own and others' body parts can be lost after brain damage, as in patients who misidentify someone else's hand as their own (pathological embodiment). Surprisingly, these patients do not use visual information to discriminate between the own and the alien hand. We asked whether this impaired visual discrimination emerges only in the ecological evaluation when the pathological embodiment is triggered by the physical alien hand (the examiner's one) or whether it emerges also when hand images are displayed on a screen. Forty right brain-damaged patients, with (E+ = 20) and without (E- = 20) pathological embodiment, and 24 healthy controls underwent two tasks in which stimuli depicting self and other hands was adopted. In the Implicit task, where participants judged which of two images matched a central target, the self-advantage (better performance with Self than Other stimuli) selectively emerges in controls, but not in patients. Moreover, E+ patients show a significantly lower performance with respect to both controls and E- patients, whereas E- patients were comparable to controls. In the Explicit task, where participants judged which stimuli belonged to themselves, both E- and E+ patients performed worst when compared to controls, but only E+ patients hyper-attributed others' hand to themselves (i.e., false alarms) as observed during the ecological evaluation. The VLSM revealed that SLF damage was significantly associated with the tendency of committing false alarm errors. We demonstrate that, in E+ patients, the ability to visually recognize the own body is lost, at both implicit and explicit level.

Reach planning with someone else's hand.

To investigate the relationship between the sense of body ownership and motor control, we capitalized on a rare bizarre disorder wherein another person's hand is misattributed to their own body, i.e., a pathological form of embodiment (E+). Importantly, despite E+ is usually associated with motor deficits, we had the opportunity to test two E+ patients with spared motor function, thus able to perform a reaching task. Crucially, these patients had proprioceptive deafferentation, allowing us to purely isolate the embodiment-dependent effect from proprioception-dependent ones that are usually associated in experimental manipulations of body ownership in healthy participants. Previous evidence suggests that the reaching movement vector is attracted towards an embodied hand during the rubber hand illusion (RHI). However, these results are confounded by the spared proprioception, whose modulation alone could explain the effects on reach planning. The neuropsychological approach employed here provides unambiguous evidence about the role of body ownership in reach planning. Indeed, three brain-damaged patients with proprioceptive deafferentation, two E+ and a well-matched control patient without pathological embodiment (E-), and 10 age-matched healthy controls underwent a reaching task wherein they had to reach for a target from a fixed starting point, while an alien hand (the co-experimenter's) was placed on the table. Irrespective of proprioception, damaged in all patients, only in E+ patients reaching errors were significantly more shifted consistently with the pathological belief, i.e., as if they planned movements from the position of the alien (embodied) hand, as compared to controls. Furthermore, with an additional experiment on healthy participants, we demonstrated that reaching errors observed during the RHI correlate with the changes in ownership. In conclusion, our neuropsychological approach suggests that when planning a reach, we do so from where our owned hand is and not from its physical location.

Wearing a Mask Shapes Interpersonal Space during COVID-19 Pandemic.

Social distancing norms have been promoted after the COVID-19 pandemic. In this work, we tested interpersonal space (IPS) in 107 subjects through a reaching-comfort distance estimation task. In the main experiment, subjects had to estimate the comfort and reach space between an avatar wearing or not wearing a face mask. We found that IPS was greater between avatars not wearing a mask with respect to stimuli with the mask on, while reaching space was not modulated. IPS increment in the NoMask condition with respect to the Mask condition correlated with anxiety traits, as shown with the State-Trait Anxiety Inventory, rather than with transient aspects related to the pandemic situation. In the control experiment, the avatars with a mask were removed to further explore the conditioning effect provided by the presence of the facial protection in the main experiment. We found a significant difference comparing this condition with the same condition of the main experiment, namely, the distances kept between avatars not wearing a mask in the main experiment were greater than those between the same stimuli in the control experiment. This showed a contextual adaptation of IPS when elements related to the actual pandemic situation were relevant. Additionally, no significant differences were found between the control experiment and the Mask condition of the main experiment, suggesting that participants had internalized social distancing norms and wearing a mask has become the new normal. Our results highlight the tendency of people in underestimating the risk of contagion when in the presence of someone wearing a mask.

Structural connectivity associated with the sense of body ownership: a diffusion tensor imaging and disconnection study in patients with bodily awareness disorder.

The brain mechanisms underlying the emergence of a normal sense of body ownership can be investigated starting from pathological conditions in which body awareness is selectively impaired. Here, we focused on pathological embodiment, a body ownership disturbance observed in brain-damaged patients who misidentify other people's limbs as their own. We investigated whether such body ownership disturbance can be classified as a disconnection syndrome, using three different approaches based on diffusion tensor imaging: (i) reconstruction of disconnectome maps in a large sample (N = 70) of stroke patients with and without pathological embodiment; (ii) probabilistic tractography, performed on the age-matched healthy controls (N = 16), to trace cortical connections potentially interrupted in patients with pathological embodiment and spared in patients without this pathological condition; (iii) probabilistic 'in vivo' tractography on two patients without and one patient with pathological embodiment. The converging results revealed the arcuate fasciculus and the third branch of the superior longitudinal fasciculus as mainly involved fibre tracts in patients showing pathological embodiment, suggesting that this condition could be related to the disconnection between frontal, parietal and temporal areas. This evidence raises the possibility of a ventral self-body recognition route including regions where visual (computed in occipito-temporal areas) and sensorimotor (stored in premotor and parietal areas) body representations are integrated, giving rise to a normal sense of body ownership.

Self-other distinction modulates the social softness illusion.

The social softness illusion (i.e., the tendency to perceive another person's skin as softer than our own) is thought to promote the sharing of social-emotional experiences because of the rewarding properties of receiving and giving social affective touch. Here we investigated whether the ability to distinguish someone else's body from our own modulates the social softness illusion. In particular, we tested whether the spatial perspective taken by the participants and seeing or not the touched arms could alter this illusion. Pairs of female participants were assigned the roles of either the giver (i.e., delivering the touches) or the receiver (i.e., being touched). We manipulated the location of the touch (palm or forearm), the spatial perspective of the receiver's body with respect to the giver's body (egocentric or allocentric perspective), and the vision of the touched body part (the giver could either see both her own and the receiver's body part, or she was blindfolded). Consistently with previous findings, the skin of another person was perceived as softer than the own one. Additionally, the illusion was present for both the forearm and the palm, and it was stronger in allocentric compared to the egocentric perspective (i.e., when the self-other distinction was clearer). These findings show that the mechanisms underpinning the ability to represent another person's body as distinct from our own modulates the social softness illusion, and thus support the role of the social softness illusion in fostering social relationships.

Monochannel Preference in Autism Spectrum Conditions Revealed by a Non-Visual Variant of Rubber Hand Illusion.

Individuals with autism spectrum conditions (ASC) are less susceptible to multisensory delusions, such as rubber hand illusion (RHI). Here, we investigate whether a monochannel variant of RHI is more effective in inducing an illusory feeling of ownership in ASC. To this aim, we exploit a non-visual variant of the RHI that, excluding vision, leverages only on the somatosensory channel. While the visual-tactile RHI does not alter the perceived hand position in ASC individuals, the tacto-tactile RHI effectively modulates proprioception to a similar extent as that found in typical development individuals. These findings suggest a more effective integration of multiple inputs originating from the same sensory channel in ASC, revealing a monochannel preference in this population.

Diametrical modulation of tactile and visual perceptual thresholds during the rubber hand illusion: a predictive coding account.

During the rubber hand illusion (RHI), the synchronous stroking of the participants' concealed hand and a visible rubber hand induces a conflict among visuo-tactile inputs, leading healthy subjects to perceive the illusion of being touched on the rubber hand, as if it were part of their body. The predictive coding theory suggests that the RHI emerges to settle the conflict, attenuating somatosensory inputs in favour of visual ones, which "capture" tactile sensations. Here, we employed the psychophysical measure of perceptual threshold to measure a behavioural correlate of the somatosensory and visual modulations, to better understand the mechanisms underpinning the illusion. Before and after the RHI, participants underwent a tactile (Experiment 1) and a visual (Experiment 2) task, wherein they had to detect stimuli slightly above the perceptual threshold. According to the predictive coding framework, we found a significant decrease of tactile detection (i.e. increased tactile perceptual threshold) and a significant increase of visual detection (i.e.  decreased visual perceptual threshold), suggesting a diametrical modulation of somatosensory and visual perceptual processes. These findings provide evidence of how our system plastically adapts to uncertainty, attributing different weights to sensory inputs to restore a coherent representation of the own body.

Behavioural evidence of altered sensory attenuation in obesity.

Body ownership (i.e., the conscious belief of owning a body) and sense of agency (i.e., being the agent of one's own movements) are part of a pre-reflective experience of bodily self, which grounds on low-level complex sensory-motor processes. Although previous literature had already investigated body ownership in obesity, sense of agency was never explored. Here, we exploited the sensory attenuation effect (i.e., an implicit marker of the sense of agency; SA effect) to investigate whether the sense of agency was altered in a sample of 18 individuals affected by obesity as compared with 18 healthy-weight individuals. In our experiment, participants were asked to rate the perceived intensity of self-generated and other-generated tactile stimuli. Healthy-weight individuals showed a significantly greater SA effect than participants affected by obesity. Indeed, while healthy-weight participants perceived self-generated stimuli as significantly less intense as compared to externally generated ones, this difference between stimuli was not reported by affected participants. Our results relative to the SA effect pinpointed an altered sense of agency in obesity. We discussed this finding within the motor control framework with reference to obesity. We encouraged future research to further explore such effect and its role in shaping the clinical features of obesity.

Seeming confines: Electrophysiological evidence of peripersonal space remapping following tool-use in humans.

The peripersonal space (PPS) is a special portion of space immediately surrounding the body, where the integration between tactile stimuli delivered on the body and auditory or visual events emanating from the environment occurs. Interestingly, PPS can widen if a tool is employed to interact with objects in the far space. However, electrophysiological evidence of such tool-use dependent plasticity in the human brain is scarce. Here, in a series of three experiments, participants were asked to respond to tactile stimuli, delivered to their right hand, either in isolation (unimodal condition) or combined with auditory stimulation, which could occur near (bimodal-near) or far from the stimulated hand (bimodal-far). According to multisensory integration spatial rule, when bimodal stimuli are presented at the same location, we expected a response enhancement (response time - RT - facilitation and event-related potential - ERP - super-additivity). In Experiment 1, we verified that RT facilitation was driven by bimodal input spatial congruency, independently from auditory stimulus intensity. In Experiment 2, we showed that our bimodal task was effective in eliciting the magnification of ERPs in bimodal conditions, with significantly larger responses in the near as compared to far condition. In Experiment 3 (main experiment), we explored tool-use driven PPS plasticity. Our audio-tactile task was performed either following tool-use (a 20-min reaching task, performed using a 145 cm-long rake) or after a control cognitive training (a 20-min visual discrimination task) performed in the far space. Following the control training, faster RTs and greater super-additive ERPs were found in bimodal-near as compared to bimodal-far condition (replicating Experiment 2 results). Crucially, this far-near differential response was significantly reduced after tool-use. Altogether our results indicate a selective effect of tool-use remapping in extending the boundaries of PPS. The present finding might be considered as an electrophysiological evidence of tool-use dependent plasticity in the human brain.

Tonic somatosensory responses and deficits of tactile awareness converge in the parietal operculum.

Although clinical neuroscience and the neuroscience of consciousness have long sought mechanistic explanations of tactile-awareness disorders, mechanistic insights are rare, mainly because of the difficulty of depicting the fine-grained neural dynamics underlying somatosensory processes. Here, we combined the stereo-EEG responses to somatosensory stimulation with the lesion mapping of patients with a tactile-awareness disorder, namely tactile extinction. Whereas stereo-EEG responses present different temporal patterns, including early/phasic and long-lasting/tonic activities, tactile-extinction lesion mapping co-localizes only with the latter. Overlaps are limited to the posterior part of the perisylvian regions, suggesting that tonic activities may play a role in sustaining tactile awareness. To assess this hypothesis further, we correlated the prevalence of tonic responses with the tactile-extinction lesion mapping, showing that they follow the same topographical gradient. Finally, in parallel with the notion that visuotactile stimulation improves detection in tactile-extinction patients, we demonstrated an enhancement of tonic responses to visuotactile stimuli, with a strong voxel-wise correlation with the lesion mapping. The combination of these results establishes tonic responses in the parietal operculum as the ideal neural correlate of tactile awareness.