The rubber hand illusion in microgravity and water immersion.

Our body has evolved in terrestrial gravity and altered gravitational conditions may affect the sense of body ownership (SBO). By means of the rubber hand illusion (RHI), we investigated the SBO during water immersion and parabolic flights, where unconventional gravity is experienced. Our results show that unconventional gravity conditions remodulate the relative weights of visual, proprioceptive, and vestibular inputs favoring vision, thus inducing an increased RHI susceptibility.

Bimanual coupling effect during a proprioceptive stimulation.

Circle-line drawing paradigm is used to study bimanual coupling. In the standard paradigm, subjects are asked to draw circles with one hand and lines with the other hand; the influence of the concomitant tasks results in two "elliptical" figures. Here we tested whether proprioceptive information evoked by muscle vibration inducing a proprioceptive illusion (PI) of movement at central level, was able to affect the contralateral hand drawing circles or lines. A multisite 80 Hz-muscle vibration paradigm was used to induce the illusion of circle- and line-drawing on the right hand of 15 healthy participants. During muscle vibration, subjects had to draw a congruent or an incongruent figure with the left hand. The ovalization induced by PI was compared with Real and Motor Imagery conditions, which already have proved to induce bimanual coupling. We showed that the ovalization of a perceived circle over a line drawing during PI was comparable to that observed in Real and Motor Imagery condition. This finding indicates that PI can induce bimanual coupling, and proprioceptive information can influence the motor programs of the contralateral hand.

Does musical interaction in a jazz duet modulate peripersonal space?

Researchers have widely studied peripersonal space (the space within reach) in the last 20 years with a focus on its plasticity following the use of tools and, more recently, social interactions. Ensemble music is a sophisticated joint action that is typically explored in its temporal rather than spatial dimensions, even within embodied approaches. We, therefore, devised a new paradigm in which two musicians could perform a jazz standard either in a cooperative (correct harmony) or uncooperative (incorrect harmony) condition, under the hypothesis that their peripersonal spaces are modulated by the interaction. We exploited a well-established audio-tactile integration task as a proxy for such a space. After the performances, we measured reaction times to tactile stimuli on the subjects' right hand and auditory stimuli delivered at two different distances, (next to the subject and next to the partner). Considering previous literature's evidence that integration of two different stimuli (e.g. a tactile and an auditory stimulus) is faster in near space compared to far space, we predicted that a cooperative interaction would have extended the peripersonal space of the musicians towards their partner, facilitating reaction times to bimodal stimuli in both spaces. Surprisingly, we obtained complementary results in terms of an increase of reaction times to tactile-auditory near stimuli, but only following the uncooperative condition. We interpret this finding as a suppression of the subject's peripersonal space or as a withdrawal from the uncooperative partner. Subjective reports and correlations between these reports and reaction times comply with that interpretation. Finally, we determined an overall better multisensory integration competence in musicians compared to non-musicians tested in the same task.

What pathological embodiment/disembodiment tell us about body representations.

In the last decade, a considerable amount of studies investigated different neuropsychological syndromes related to the disorder of body awareness. In this paper, we shall review neuropsychological evidence of pathological embodiment/disembodiment conditions with the aim of describing the major common features, and the complementary characteristics, that may suggest the structure and function of a shared body representation. In particular, we shall first discuss experimental studies conducted on a bizarre disorder of body ownership we recently described [i.e., brain-damaged patients claiming that another person's hand belongs to them (Pathological Embodiment, PE)]. Then complementary syndromes, with an apparent opposite attitude with respect to the PE (i.e., somatoparaphrenia and xenomelia, which is part of the Body Integrity Identity Disorders) will be also considered. We shall discuss the behavioural similarities/differences between these complementary disturbances also referring to already existing conceptual knowledge and proposals about body representation.

Effect of tool-use observation on metric body representation and peripersonal space.

In everyday life, we constantly act and interact with objects and with others' people through our body. To properly perform actions, the representations of the dimension of body-parts (metric body representation, BR) and of the space surrounding the body (peripersonal space, PPS) need to be constantly updated. Previous evidence has shown that BR and PPS representation are highly flexible, being modulated by sensorimotor experiences, such as the active use of tools to reach objects in the far space. In this study, we investigate whether the observation of another person using a tool to interact with objects located in the far space is sufficient to influence the plasticity of BR and PPS representation in a similar way to active tool-use. With this aim, two groups of young healthy participants were asked to perform 20 min trainings based on the active use of a tool to retrieve far cubes (active tool-use) and on the first-person observation of an experimenter doing the same tool-use training (observational tool-use). Behavioural tasks adapted from literature were used to evaluate the effects of the active and observational tool-use on BR (body-landmarks localization task-group 1), and PPS (audio-tactile interaction task - group 2). Results show that after active tool-use, participants perceived the length of their arm as longer than at baseline, while no significant differences appear after observation. Similarly, significant modifications in PPS representation, with comparable multisensory facilitation on tactile responses due to near and far sounds, were seen only after active tool-use, while this did not occur after observation. Together these results suggest that a mere observational training could not be sufficient to significantly modulate BR or PPS. The dissociation found in the active and observational tool-use points out differences between action execution and action observation, by suggesting a fundamental role of the motor planning, the motor intention, and the related sensorimotor feedback in driving BR and PPS plasticity.

Immersive virtual reality reveals that visuo-proprioceptive discrepancy enlarges the hand-centred peripersonal space.

Vision and proprioception, informing the system about the body position in space, seem crucial in defining the boundary of the peripersonal space (PPS). What happens to the PPS representation when a conflict between vision and proprioception arises? We capitalize on the Immersive Virtual Reality to dissociate vision and proprioception by presenting the participants' 3D hand image in congruent/incongruent positions with respect to the participants' real hand. To measure the hand-centred PPS, we exploit multisensory integration occurring when visual stimuli are delivered simultaneously with tactile stimuli applied to a body district; i.e., visual enhancement of touch (VET). Participants are instructed to respond to tactile stimuli while ignoring visual stimuli (red LED), which can appear either near to or far from the hand receiving tactile (electrical) stimuli. The results show that, when vision and proprioception are congruent (i.e., real and virtual hand coincide), a space-dependent modulation of the VET effect occurs (with faster responses when visual stimuli are near to than far from the stimulated hand). Contrarily, when vision and proprioception are incongruent (i.e., a discrepancy between real and virtual hand is present), a comparable VET effect is observed when visual stimuli occur near to the real hand and when they occur far from it, but close to the virtual hand. These findings, also confirmed by the independent estimate of a Bayesian Causal Inference model, suggest that, when the visuo-proprioceptive discrepancy makes the coding of the hand position less precise, the hand-centred PPS is enlarged, likely to optimize reactions to external events.

Agent-dependent modulation of corticospinal excitability during painful transcutaneous electrical stimulation.

Pain has an inhibitory effect on the corticospinal excitability that has been interpreted as an evolutionary mechanism, directed to down-regulate cortical activity in order to facilitate rapid protective spinal reflexes. Here, we focused on the link between defensive mechanisms and motor system and we asked whether voluntary actions can modulate the corticospinal excitability during painful stimulations. To this aim, we manipulated the volition-related aspects of our paradigm by comparing conditions in which either the participant (self-generated action) or the experimenter (other-generated action) pressed the button to deliver painful high-intensity transcutaneous electric shocks to the right digit V. MEPs to TMS were recorded from the FDI and APB muscles of the stimulated hand. A compelling agent-dependent modulation of the corticospinal excitability was found, showing, in self-generated compared to other-generated actions, a significantly lower inhibitory effect, as measured by greater MEP amplitude. This finding suggests a top-down modulation of volitional actions on defensive mechanisms, promoting the view that predictive information from the motor system attenuates the responses to the foreseeable adverse events generated by one's own actions as compared to unpredictable events generated by someone else's actions.

The sense of body-ownership gates cross-modal improvement of tactile extinction in brain-damaged patients.

When body ownership is impaired after brain-damage, the capacity to discriminate between one's own and others' body-parts is lost. Delusional body-ownership has been recently described in patients who misidentify someone else's limb as their own (pathological embodiment) whenever it is positioned in a body congruent position. This delusion can be frequently associated with somatosensory and attentional deficits. Here, we leveraged the phenomenon of tactile extinction, as this clinical sign can be substantially ameliorated when contralesional touches are combined with proximal visual stimulation. Is body ownership a necessary prerequisite to modulate cross-modal processing and thus reducing tactile extinction? Fourteen patients with tactile extinction (TE+) took part in the study: eight of them with pathological embodiment (E+, experimental group) and six of them without pathological embodiment (E-, control group). In two different paradigms, differing for the nature of visuo-tactile stimuli, bilateral tactile stimulation of the patients' hands was combined with visual stimuli occurring on A) their own contralesional (affected) hand, B) the examiner's hand (embodied in E+), or C) a neutral object. In both groups, visual stimuli proximal to the own hand significantly improved contralesional tactile detection, while visual stimuli occurring on the neutral object did not. Crucially, only in E+TE+ patients did visual stimuli on the examiner's (embodied) hand improve contralesional tactile detection. This finding shows that cross-modal visuo-tactile integration is conditional to body-ownership, so that it ameliorates tactile extinction when visual stimuli occur on what is believed to be one's own body. From a clinical point of view, this study suggests that the effectiveness of cross-modal rehabilitative intervention can benefit from a careful evaluation of the patients' sense of body-ownership, so often impaired after brain-damage.

Aesthetic appreciation of musical intervals enhances behavioural and neurophysiological indexes of attentional engagement and motor inhibition.

From Kant to current perspectives in neuroaesthetics, the experience of beauty has been described as disinterested, i.e. focusing on the stimulus perceptual features while neglecting self-referred concerns. At a neurophysiological level, some indirect evidence suggests that disinterested aesthetic appreciation might be associated with attentional enhancement and inhibition of motor behaviour. To test this hypothesis, we performed three auditory-evoked potential experiments, employing consonant and dissonant two-note musical intervals. Twenty-two volunteers judged the beauty of intervals (Aesthetic Judgement task) or responded to them as fast as possible (Detection task). In a third Go-NoGo task, a different group of twenty-two participants had to refrain from responding when hearing intervals. Individual aesthetic judgements positively correlated with response times in the Detection task, with slower motor responses for more appreciated intervals. Electrophysiological indexes of attentional engagement (N1/P2) and motor inhibition (N2/P3) were enhanced for more appreciated intervals. These findings represent the first experimental evidence confirming the disinterested interest hypothesis and may have important applications in research areas studying the effects of stimulus features on learning and motor behaviour.

Bodily ownership modulation in defensive responses: physiological evidence in brain-damaged patients with pathological embodiment of other's body parts.

Do conscious beliefs about the body affect defensive mechanisms within the body? To answer this question we took advantage from a monothematic delusion of bodily ownership, in which brain-damaged patients misidentify alien limbs as their own. We investigated whether the delusional belief that an alien hand is their own hand modulates a subcortical defensive response, such as the hand-blink reflex. The blink, dramatically increases when the threated hand is inside the defensive peripersonal-space of the face. In our between-subjects design, including patients and controls, the threat was brought near the face either by the own hand or by another person's hand. Our results show an ownership-dependent modulation of the defensive response. In controls, as well as in the patients' intact-side, the response enhancement is significantly greater when the threat was brought near the face by the own than by the alien hand. Crucially, in the patients' affected-side (where the pathological embodiment occurs), the alien (embodied) hand elicited a response enhancement comparable to that found when the threat is brought near the face by the real hand. These findings suggest the existence of a mutual interaction between our conscious beliefs about the body and the physiological mechanisms within the body.

Interpersonal interactions and empathy modulate perception of threat and defensive responses.

The defensive peripersonal space (DPPS) is a vital "safety margin" surrounding the body. When a threatening stimulus is delivered inside the DPPS, subcortical defensive responses like the hand-blink reflex (HBR) are adjusted depending on the perceived threat content. In three experiments, we explored whether and how defensive responses are affected by the interpersonal interaction within the DPPS of the face. In Experiment 1, we found that the HBR is enhanced when the threat is brought close to the face not only by one's own stimulated hand, but also by another person's hand, although to a significantly lesser extent. In Experiments 2 and 3, we found that the HBR is also enhanced when the hand of the participant enters the DPPS of another individual, either in egocentric or in allocentric perspective. This enhancement is larger in participants with strong empathic tendency when the other individual is in a third person perspective. These results indicate that interpersonal interactions shape perception of threat and defensive responses. These effects are particularly evident in individuals with greater tendency to having empathic concern to other people.

Bimanual non-congruent actions in motor neglect syndrome: a combined behavioral/fMRI study.

In Motor Neglect (MN) syndrome, a specific impairment in non-congruent bimanual movements has been described. In the present case-control study, we investigated the neuro-functional correlates of this behavioral deficit. Two right-brain-damaged (RBD) patients, one with (MN+) and one without (MN-) MN, were evaluated by means of functional Magnetic Resonance Imaging (fMRI) in a bimanual Circles-Lines (CL) paradigm. Patients were requested to perform right-hand movements (lines-drawing) and, simultaneously, congruent (lines-drawing) or non-congruent (circles-drawing) left-hand movements. In the behavioral task, MN- patient showed a bimanual-coupling-effect, while MN+ patient did not. The fMRI study showed that in MN-, a fronto-parietal network, mainly involving the pre-supplementary motor area (pre-SMA) and the posterior parietal cortex (PPC), was significantly more active in non-congruent than in congruent conditions, as previously shown in healthy subjects. On the contrary, MN+ patient showed an opposite pattern of activation both in pre-SMA and in PPC. Within this fronto-parietal network, the pre-SMA is supposed to exert an inhibitory influence on the default coupling of homologous muscles, thus allowing the execution of non-congruent movements. In MN syndrome, the described abnormal pre-SMA activity supports the hypothesis that a failure to inhibit ipsilesional motor programs might determine a specific impairment of non-congruent movements.