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.

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.

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.

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.

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.

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.

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.

Face-like configurations modulate electrophysiological mismatch responses.

The human face is one of the most salient stimuli in the environment. It has been suggested that even basic face-like configurations (three dots composing a downward pointing triangle) may convey salience. Interestingly, stimulus salience can be signaled by mismatch detection phenomena, characterized by greater amplitudes of event-related potentials (ERPs) in response to relevant novel stimulation as compared to non-relevant repeated events. Here, we investigate whether basic face-like stimuli are salient enough to modulate mismatch detection phenomena. ERPs are elicited by a pair of sequentially presented visual stimuli (S1-S2), delivered at a constant 1-s interval, representing either a face-like stimulus (Upright configuration) or three neutral configurations (Inverted, Leftwards, and Rightwards configurations), that are obtained by rotating the Upright configuration along the three different axes. In pairs including a canonical face-like stimulus, we observe a more effective mismatch detection mechanism, with significantly larger N270 and P300 components when S2 is different from S1 as compared to when S2 is identical to S1. This ERP modulation, not significant in pairs excluding face-like stimuli, reveals that mismatch detection phenomena are significantly affected by basic face-like configurations. Even though further experiments are needed to ascertain whether this effect is specifically elicited by face-like configuration rather than by particular orientation changes, our findings suggest that face essential, structural attributes are salient enough to affect change detection processes.

Like the back of my hand: Visual ERPs reveal a specific change detection mechanism for the bodily self.

The ability to identify our own body is considered a pivotal marker of self-awareness. Previous research demonstrated that subjects are more efficient in the recognition of images representing self rather than others' body effectors (self-advantage). Here, we verified whether, at an electrophysiological level, bodily-self recognition modulates change detection responses. In a first EEG experiment (discovery sample), event-related potentials (ERPs) were elicited by a pair of sequentially presented visual stimuli (vS1; vS2), representing either the self-hand or other people's hands. In a second EEG experiment (replicating sample), together with the previously described visual stimuli, also a familiar hand was presented. Participants were asked to decide whether vS2 was identical or different from vS1. Accuracy and response times were collected. In both experiments, results confirmed the presence of the self-advantage: participants responded faster and more accurately when the self-hand was presented. ERP results paralleled behavioral findings. Anytime the self-hand was presented, we observed significant change detection responses, with a larger N270 component for vS2 different rather than identical to vS1. Conversely, when the self-hand was not included, and even in response to the familiar hand in Experiment 2, we did not find any significant modulation of the change detection responses. Overall our findings, showing behavioral self-advantage and the selective modulation of N270 for the self-hand, support the existence of a specific mechanism devoted to bodily-self recognition, likely relying on the multimodal (visual and sensorimotor) dimension of the bodily-self representation. We propose that such a multimodal self-representation may activate the salience network, boosting change detection effects specifically for the self-hand.