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.

Modulation of vestibular input by short-term head-down bed rest affects somatosensory perception: implications for space missions.

Introduction: On Earth, self-produced somatosensory stimuli are typically perceived as less intense than externally generated stimuli of the same intensity, a phenomenon referred to as somatosensory attenuation (SA). Although this phenomenon arises from the integration of multisensory signals, the specific contribution of the vestibular system and the sense of gravity to somatosensory cognition underlying distinction between self-generated and externally generated sensations remains largely unknown. Here, we investigated whether temporary modulation of the gravitational input by head-down tilt bed rest (HDBR)-a well-known Earth-based analog of microgravity-might significantly affect somatosensory perception of self- and externally generated stimuli. Methods: In this study, 40 healthy participants were tested using short-term HDBR. Participants received a total of 40 non-painful self- and others generated electrical stimuli (20 self- and 20 other-generated stimuli) in an upright and HDBR position while blindfolded. After each stimulus, they were asked to rate the perceived intensity of the stimulation on a Likert scale. Results: Somatosensory stimulations were perceived as significantly less intense during HDBR compared to upright position, regardless of the agent administering the stimulus. In addition, the magnitude of SA in upright position was negatively correlated with the participants' somatosensory threshold. Based on the direction of SA in the upright position, participants were divided in two subgroups. In the subgroup experiencing SA, the intensity rating of stimulations generated by others decreased significantly during HDBR, leading to the disappearance of the phenomenon of SA. In the second subgroup, on the other hand, reversed SA was not affected by HDBR. Conclusion: Modulation of the gravitational input by HDBR produced underestimation of somatosensory stimuli. Furthermore, in participants experiencing SA, the reduction of vestibular inputs by HDBR led to the disappearance of the SA phenomenon. These findings provide new insights into the role of the gravitational input in somatosensory perception and have important implications for astronauts who are exposed to weightlessness during space missions.

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.

When Right Goes Left: Phantom Touch Induced by Mirror Box Procedure in Healthy Individuals.

In the present article, we investigated the possibility of inducing phantom tactile sensations in healthy individuals similar to those that we observed in patients after stroke. On the basis of previous research, we assumed that manipulating visual feedbacks may guide and influence, under certain conditions, the phenomenal experience of touch. To this aim, we used the Tactile Quadrant Stimulation (TQS) test in which subjects, in the crucial condition, must indicate whether and where they perceive a double tactile stimulation applied simultaneously in different quadrants of the two hands (asymmetrical Double Simultaneous Stimulation trial, Asym-DSS). The task was performed with the left-hand out of sight and the right-hand reflected in a mirror so that the right-hand reflected in the mirror looks like the own left-hand. We found that in the Asym-DSS trial, the vision of the right-hand reflected in the mirror and stimulated by a tactile stimulus elicited on the left-hand the sensation of having been touched in the same quadrant as the right-hand. In other words, we found in healthy subjects the same phantom touch effect that we previously found in patients. We interpreted these results as modulation of tactile representation by bottom-up (multisensory integration of stimuli coming from the right real and the right reflected hand) and possibly top-down (body ownership distortion) processing triggered by our experimental setup, unveiling bilateral representation of touch.

From embodying tool to embodying alien limb: sensory-motor modulation of personal and extrapersonal space.

Years ago, it was demonstrated (e.g., Rizzolatti et al. in Handbook of neuropsychology, Elsevier Science, Amsterdam, 2000) that the brain does not encode the space around us in a homogeneous way, but through neural circuits that map the space relative to the distance that objects of interest have from the body. In monkeys, relatively discrete neural systems, characterized by neurons with specific neurophysiological responses, seem to be dedicated either to represent the space that can be reached by the hand (near/peripersonal space) or to the distant space (far/extrapersonal space). It was also shown that the encoding of spaces has dynamic aspects because they can be remapped by the use of tools that trigger different actions (e.g., Iriki et al. 1998). In this latter case, the effect of the tool depends on the modulation of personal space, that is the space of our body. In this paper, I will review and discuss selected research, which demonstrated that also in humans: 1 spaces are encoded in a dynamic way; 2 encoding can be modulated by the use of tool that the system comes to consider as parts of the own body; 3 body representations are not fixed, but they are fragile and subject to change to the point that we can incorporate not only the tools necessary for action, but even limbs belonging to other people. What embodiment of tools and of alien limb tell us about body representations is then briefly discussed.

Zero gravity induced by parabolic flight enhances automatic capture and weakens voluntary maintenance of visuospatial attention.

Orienting attention in the space around us is a fundamental prerequisite for willed actions. On Earth, at 1 g, orienting attention requires the integration of vestibular signals and vision, although the specific vestibular contribution to voluntary and automatic components of visuospatial attention remains largely unknown. Here, we show that unweighting of the otolith organ in zero gravity during parabolic flight, selectively enhances stimulus-driven capture of automatic visuospatial attention, while weakening voluntary maintenance of covert attention. These findings, besides advancing our comprehension of the basic influence of the vestibular function on voluntary and automatic components of visuospatial attention, may have operational implications for the identification of effective countermeasures to be applied in forthcoming human deep space exploration and habitation, and on Earth, for patients' rehabilitation.

The anatomo-clinical picture of the pathological embodiment over someone else's body part after stroke.

Recently, a monothematic delusion of body ownership due to brain damage (i.e., the embodiment of someone else's body part within the patient's sensorimotor system) has been extensively investigated. Here we aimed at defining in-depth the clinical features and the neural correlates of the delusion. Ninety-six stroke patients in a sub-acute or chronic phase of the illness were assessed with a full ad-hoc protocol to evaluate the embodiment of an alien arm under different conditions. A sub-group of seventy-five hemiplegic patients was also evaluated for the embodiment of the movements of the alien arm. Fifty-five patients were studied to identify the neural bases of the delusion by means of voxel-based lesion-symptom mapping approach. Our results show that, in forty percent of the whole sample, simply viewing the alien arm triggered the delusion, but only if it was a real human arm and that was seen from a 1st person perspective in an anatomically-correct position. In the hemiplegic sub-group, the presence of the embodiment of the alien arm was always accompanied by the embodiment of its passive and active movements. Furthermore, the delusion was significantly associated to primary proprioceptive deficits and to damages of the corona radiata and the superior longitudinal fasciculus. To conclude, we show that the pathological embodiment of an alien arm is well-characterized by recurrent and specific features and might be explained as a disconnection deficit, mainly involving white matter tracts. The proposed exhaustive protocol can be successfully employed to assess stroke-induced disorders of body awareness, unveiling even their more undetectable or covert clinical forms.

Direct electrical stimulation of the premotor cortex shuts down awareness of voluntary actions.

A challenge for neuroscience is to understand the conscious and unconscious processes underlying construction of willed actions. We investigated the neural substrate of human motor awareness during awake brain surgery. In a first experiment, awake patients performed a voluntary hand motor task and verbally monitored their real-time performance, while different brain areas were transiently impaired by direct electrical stimulation (DES). In a second experiment, awake patients retrospectively reported their motor performance after DES. Based on anatomo-clinical evidence from motor awareness disorders following brain damage, the premotor cortex (PMC) was selected as a target area and the primary somatosensory cortex (S1) as a control area. In both experiments, DES on both PMC and S1 interrupted movement execution, but only DES on PMC dramatically altered the patients' motor awareness, making them unconscious of the motor arrest. These findings endorse PMC as a crucial hub in the anatomo-functional network of human motor awareness.

Acute and cumulative effects of rTMS on behavioural and EMG parameters in Focal Hand Dystonia.

Previous studies suggest that low-frequency repetitive Transcranial Magnetic Stimulation (rTMS) over contralateral premotor cortex (PMC) might ameliorate Focal Hand Dystonia (FHD) symptoms. In the present study behavioral and muscle activity outcomes were explored in a patient with FHD following a single and multiple sessions of rTMS. The patient's behavior was assessed on handwriting tasks, while surface EMG signals were recorded. In Experiment 1 evaluations were performed before and after one session of active and sham 1Hz rTMS over contralateral PMC. In Experiment 2, evaluations were performed before and after six sessions of the same treatment. In Experiment 1 active rTMS improved the patient's performance, although the EMG amplitude did not change. In Experiment 2, the patient showed an improvement of performance along with a decrease of 20% in the EMG amplitude. These results demonstrated that a single session of rTMS ameliorated the patient's performance, while multiple sessions were necessary to reduce muscles activity.

Phantom touch: How to unmask sensory unawareness after stroke.

Comprehending the nature of tactile disorders following brain damage is crucial to understand how the brain constructs sensory awareness. Stroke patients may be unaware of being touched on the affected hand if, simultaneously, they are touched on the unaffected hand (i.e., tactile extinction). More rarely, they feel touches on the two hands, when they are solely touched on the unaffected hand (i.e., synchiria). Using a novel assessment tool, we investigated whether in stroke patients with apparent intact tactile awareness on standard evaluation, tactile extinction might be possibly masked by phantom (synchiric) sensations (i.e., elicited by ipsilesional stimulation) arising exclusively during Double Simultaneous Stimulation (DSS). Patients with right (n = 17) and left (n = 8) hemisphere lesions and age-matched healthy controls (n = 13) were tested with the Tactile Quadrant Stimulation test, consisting in delivering unilateral or bilateral touches to one of four quadrants, identified on the participants' hands. In DSS trials, stimuli were applied to asymmetric quadrants. Participants reported the side(s) and then pointed to the site(s) of stimulation. We found that, with the exception of one patient who showed tactile extinction, about 50% of patients with overall intact tactile perception on classical evaluation, although reporting two stimuli in DSS, failed in pointing to the correct contralesional stimulated site. They reported the felt sensation in positions that corresponded to the ipsilesional stimulated sites. Thus apparent detections of contralesional touches in DSS were accounted for by 'phantom' sensations of ipsilesional stimulation that masked unawareness of contralesional touches when classic assessment was applied. Preliminary lesion analyses indicate that the symptom was associated with damage to structures often affected in tactile extinction. These findings, while unveiling important underestimation of the patients' neurological condition, provide a framework for investigating bihemispheric contributions to altered tactile perception following stroke.

Transcranial Magnetic Stimulation of Posterior Parietal Cortex Modulates Line-Length Estimation but Not Illusory Depth Perception.

Transcranial Magnetic Stimulation (TMS) may affect attentional processing when applied to the right posterior parietal cortex (PPC) of healthy participants in line with neuropsychological and neuroimaging evidence on the neural bases of this cognitive function. Specifically, the application of TMS to right PPC induces a rightward attentional bias on line length estimation in healthy participants (i.e., neglect-like bias), mimicking the rightward bias shown by patients with unilateral spatial neglect after damage of the right PPC. With the present study, we investigated whether right PPC might play a crucial role in attentional processing of illusory depth perception, given the evidence that a rightward bias may be observed in patients with neglect during perception of the Necker Cube (NC). To this end, we investigated the effects of low-frequency rTMS applied to the right or left PPC on attentional disambiguation of the NC in two groups of healthy participants. To control for the effectiveness of TMS on visuospatial attention, rTMS effects were also assessed on a frequently used line length estimation (i.e., the Landmark Task or LT). Both groups also received sham stimulation. RTMS of the right or left PPC did not affect NC perception. On the other hand, rTMS of the right PPC (but not left PPC) induces neglect-like bias on the LT, in line with previous studies. These findings confirm that right PPC is involved in deployment of spatial attention on line length estimation. Interestingly, they suggest that this brain region does not critically contribute to deployment of visuospatial attention during attentional disambiguation of the Necker Cube. Future investigations, targeting different areas of fronto-parietal circuits, are necessary to further explore the neuro-functional bases of attentional contribution to illusory depth perception.

Shared neurocognitive mechanisms of attenuating self-touch and illusory self-touch.

Despite the fact that any successful achievement of willed actions necessarily entails the sense of body ownership (the feeling of owning the moving body parts), it is still unclear how this happens. To address this issue at both behavioral and neural levels, we capitalized on sensory attenuation (SA) phenomenon (a self-generated stimulus is perceived as less intense than an identical externally generated stimulus). We compared the intensity of somatosensory stimuli produced by one's own intended movements and by movements of an embodied fake hand. Then, we investigated if in these two conditions SA was equally affected by interfering with the activity of the supplementary motor area (SMA; known to be related to motor intention and SA) using single-pulse transcranial magnetic stimulation. We showed that ownership of the fake hand triggered attenuation of somatosensory stimuli generated by its movements that were comparable to the attenuation of self-generated stimuli. Furthermore, disrupting the SMA eliminated the SA effect regardless of whether it was triggered by actual participant's movements or by illusory ownership. Our findings suggest that SA triggered by body ownership relies, at least in part, on the activation of the same brain structures as SA triggered by motor-related signals.

To Move or Not to Move? Functional Role of Ventral Premotor Cortex in Motor Monitoring During Limb Immobilization.

Anatomo-clinical evidence from motor-awareness disorders after brain-damages suggests that the premotor cortex (PMC) is involved in motor-monitoring of voluntary actions. Indeed, PMC lesions prevent patients from detecting the mismatch between intended, but not executed, movements with the paralyzed limb. This functional magnetic resonance imaging study compared, in healthy subjects, free movements against blocked movements, precluded by a cast. Cast-related corticospinal excitability changes were investigated by using transcranial magnetic stimulation. Immediately after the immobilization, when the cast prevented the execution of left-hand movements, the contralateral right (ventral) vPMC showed both increased hemodynamic activity and increased functional connectivity with the hand area in the right somatosensory cortex, suggesting a vPMC involvement in detecting the mismatch between planned and executed movements. Crucially, after 1 week of immobilization, when the motor system had likely learned that no movement could be executed and, therefore, predictions about motor consequences were changed, vPMC did not show the enhanced activity as if no incongruence has to be detected. This can be interpreted as a consequence of the plastic changes induced by long-lasting immobilization, as also proved by the cast-related corticospinal excitability modulation in our subjects. The present findings highlight the crucial role of vPMC in the anatomo-functional network generating the human motor-awareness.

Dissociation between executed and imagined bimanual movements in autism spectrum conditions.

Autism spectrum conditions (ASC) are characterized by social-communicative deficits and repetitive stereotyped behaviors. Altered motor coordination is also observed and a dysfunction of motor imagery has been recently reported on implicit tasks. However, no information on explicit motor imagery abilities is available in ASC. Here, we employed a spatial bimanual task to concurrently assess motor coordination and explicit motor imagery in autism. A secondary objective of the study was to evaluate these abilities across two populations of ASC, namely adolescents and adults with ASC. To this aim, we took advantage of the circles-lines task in which where participants were asked to continuously draw: right hand lines (unimanual condition); right hand lines and left hand circles (bimanual condition); right hand lines while imagining to draw left hand circles (imagery condition). For each participant, an Ovalization Index (OI) was calculated as a deviation of the right hand drawing trajectory from an absolute vertical axis. Results showed a significant and similar coupling effect in the bimanual condition (i.e., a significant increase of the OI values with respect to the unimanual condition) in both controls and ASC participants. On the contrary, in the imagery condition, a significant coupling effect was found only in controls. Furthermore, adult controls showed a significantly higher imagery coupling effect in comparison to all the other groups. These results demonstrate that atypical motor imagery processes in ASC are not limited to implicit tasks and suggest that development of neural structures involved in motor imagery are immature in ASC. Autism Res 2018, 11: 376-384. © 2017 International Society for Autism Research, Wiley Periodicals, Inc. LAY SUMMARY: Autism spectrum conditions (ASC) are characterized by social-communicative and motor coordination difficulties but in many cases also by an impaired capability to imagine movements. In this study we found that while two handed coordination in ASC can be developed as well as in typically developed persons, the development of motor imagery could still be immature in ASC, leading to difficulties in imagining, understanding as well as programming and coordinating complex movements.

Feeling touch on the own hand restores the capacity to visually discriminate it from someone else' hand: Pathological embodiment receding in brain-damaged patients.

The sense of body ownership, i.e., the belief that a specific body part belongs to us, can be selectively impaired in brain-damaged patients. Recently, a pathological form of embodiment has been described in patients who, when the examiner's hand is located in a body-congruent position, systematically claim that it is their own hand (E+ patients). This paradoxical behavior suggests that, in these patients, the altered sense of body ownership also affects their capacity of visually discriminating the body-identity details of the own and the alien hand, even when both hands are clearly visible on the table. Here, we investigated whether, in E+ patients with spared tactile sensibility, a coherent body ownership could be restored by introducing a multisensory conflict between what the patients feel on the own hand and what they see on the alien hand. To this aim, we asked the patients to rate their sense of body ownership over the alien hand, either after segregated tactile stimulations of the own hand (out of view) and of the alien hand (visible) or after synchronous and asynchronous tactile stimulations of both hands, as in the rubber hand illusion set-up. Our results show that, when the tactile sensation perceived on the patient's own hand was in conflict with visual stimuli observed on the examiner's hand, E+ patients noticed the conflict and spontaneously described visual details of the (visible) examiner's hand (e.g., the fingers length, the nails shape, the skin color…), to conclude that it was not their own hand. These data represent the first evidence that, in E+ patients, an incongruent visual-tactile stimulation of the own and of the alien hand reduces, at least transitorily, the delusional body ownership over the alien hand, by restoring the access to the perceptual self-identity system, where visual body identity details are stored.

Movements and body ownership: Evidence from the rubber hand illusion after mechanical limb immobilization.

There is no consensus on whether, and to what extent, actions contribute to constructing awareness of one's own body. Here we investigated at both physiological and behavioral level whether a prolonged limb immobilization affects body ownership. We tested a group of healthy participants, whose left-hand movements were prevented by a cast for one week, and a control group without any movement restriction. In both groups, we measured the strength of the rubber hand illusion (i.e., proprioceptive shift and questionnaire on ownership) and the physiological parameters known to be modulated by short-term arm immobilization (i.e., resting motor threshold, motor evoked potentials and force parameters) before and after the week of immobilization. Our results showed stronger illusory effects on the immobilized hand on both behavioral indexes and weaker illusory effects on the non-immobilized hand on the questionnaire. Additionally, the increased proprioceptive shift was positively correlated to the motor threshold of the contralateral hemisphere. Our findings show at both behavioral and physiological level that altering those movement-related signals which constantly stem from our own body parts, modulates the experience of those body parts as mine. This, in turn, supports the view of a direct role of actions in the developing and maintaining a coherent body ownership.

The role of premotor and parietal cortex during monitoring of involuntary movement: A combined TMS and tDCS study.

Converging evidence on voluntary actions underlays the existence of a motor monitoring system able to compare the predicted and the actual consequences of our movements. In this context, both the premotor cortex (PMC) and the posterior parietal cortex (PPC) play a role in action monitoring and awareness. The present study explores the role of PMC and PPC in monitoring involuntary muscle contractions induced by transcranial magnetic stimulation (TMS) over the hand motor area. To this aim, the effects of transcranial direct current stimulation (tDCS) over PMC and PPC were examined. Thirty-six healthy subjects were asked to perform a motor monitoring task (i.e., to verbally report hand twitches induced by TMS) after 10 min of tDCS. Through three experiments, the effects of cathodal, anodal and sham tDCS over the left and the right hemispheres were compared. Our results show that cathodal tDCS over the right PMC does not affect the monitoring of involuntary movements. By contrast, tDCS over both the right and the left PPC affects motor monitoring, depending on the current polarity: while cathodal tDCS increases the feeling of phantom-like movements (which actually did not occur), anodal tDCS impairs the ability to detect involuntary hand twitches (which actually took place). These findings show that the PMC is not involved in motor monitoring of involuntary movements; rather, the PPC, where multisensory stimuli converge and are processed, seems to play a crucial role.

Adolescents' and Young Adults' Naïve Understandings of the Economic Crisis.

Over the last decade, Financial Literacy (FL) and interventions aimed at improving it, that is Financial Education (FE), have been the focus of increased attention from economists, governments, and international organizations such as the world Bank and OECD, but much less by scholars in the fields of Learning and Instruction. We examined open-ended written answers on the causes of the economic crisis that started in 2007-2008, as given by 381 Italian secondary school and university students, and 268 Swiss Italian-speaking secondary school students. Most Italian students mentioned internal political causes (i.e., corrupt politicians or inefficiency of the government), whereas Swiss students mentioned banks more often. International factors were rarely mentioned by either group, and explanations were generally very poor, listing a few causes without making connections between them. These findings indicate the need for economics education aimed at making people more knowledgeable of the workings of the economic system and the effects of financial systems on the real economy.

Sensing the body, representing the body: Evidence from a neurologically based delusion of body ownership.

Humans experience their own body as unitary and monolithic in nature. However, recent findings in cognitive neuroscience seem to suggest that body awareness has a complex and multifaceted structure that can be dissociated in several subcomponents, possibly underpinned by different brain circuits. In the present paper, we focus on a recently reported neuropsychological disorder of body ownership in which patients misattribute to themselves someone else's arm and its movements. As first, we briefly review the clinical and functional features of this disorder. Secondly, we attempt to explain the nature of the delusion and to gain new hints regarding the mechanisms subserving the construction and the maintenance of the sense of body ownership in the intact brain functioning.

Effects of attentional and cognitive variables on unilateral spatial neglect.

Patients with visuospatial neglect when asked to cancel targets partially or totally omit to cancel contralesional stimuli. It has been shown that increasing the attentional demands of the cancellation task aggravates neglect contralesionally. However, some preliminary evidence also suggests that neglect might be worsened by engaging the patient in a demanding, non-spatial, cognitive activity (i.e. a mathematical task). We studied cancellation performance of 16 patients with right-hemisphere lesions, 8 with neglect, 8 without neglect, and 8 age-matched healthy control participants by means of five cancellation tasks which varied for the degree of attentional and/or high level cognitive demands (preattentive and attentive search of a visual target, searching for numbers containing the digit 3, even numbers, and multiples of 3). Results showed that attentive search of visual targets, relative to the preattentive search condition, aggravated neglect patients' performance. Moreover, searching for multiples not only worsened spatial neglect contralesionally, but also slowed down performance of patients with right-hemisphere lesions without neglect. Our findings further demonstrate the presence of specific deficits of attention in neglect. In addition, the worse performance of patients without neglect in the 'multiples of 3' task is consistent with the evidence that right-hemisphere lesions per se impair the ability to maintain attention (i.e. sustained attention). This suggests that the exacerbation of neglect during execution of a demanding, non-spatial, cognitive task might be explained by a deficit of sustained attention in addition to a selective deficit of spatial attention.