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BACKGROUND: Sensory reafferents are crucial to correct our posture and movements, both reflexively and in a cognitively driven manner. They are also integral to developing and maintaining a sense of agency for our actions. In cases of compromised reafferents, such as for persons with amputated or congenitally missing limbs, or diseases of the peripheral and central nervous systems, augmented sensory feedback therefore has the potential for a strong, neurorehabilitative impact. We here developed an untethered vibrotactile garment that provides walking-related sensory feedback remapped non-invasively to the wearer's back. Using the so-called FeetBack system, we investigated if healthy individuals perceive synchronous remapped feedback as corresponding to their own movement (motor awareness) and how temporal delays in tactile locomotor feedback affect both motor awareness and walking characteristics (adaptation). METHODS: We designed the system to remap somatosensory information from the foot-soles of healthy participants (N = 29), using vibrotactile apparent movement, to two linear arrays of vibrators mounted ipsilaterally on the back. This mimics the translation of the centre-of-mass over each foot during stance-phase. The intervention included trials with real-time or delayed feedback, resulting in a total of 120 trials and approximately 750 step-cycles, i.e. 1500 steps, per participant. Based on previous work, experimental delays ranged from 0ms to 1500ms to include up to a full step-cycle (baseline stride-time: µ = 1144 ± 9ms, range 986-1379ms). After each trial participants were asked to report their motor awareness. RESULTS: Participants reported high correspondence between their movement and the remapped feedback for real-time trials (85 ± 3%, µ ± σ), and lowest correspondence for trials with left-right reversed feedback (22 ± 6% at 600ms delay). Participants further reported high correspondence of trials delayed by a full gait-cycle (78 ± 4% at 1200ms delay), such that the modulation of motor awareness is best expressed as a sinusoidal relationship reflecting the phase-shifts between actual and remapped tactile feedback (cos model: 38% reduction of residual sum of squares (RSS) compared to linear fit, p < 0.001). The temporal delay systematically but only moderately modulated participant stride-time in a sinusoidal fashion (3% reduction of RSS compared a linear fit, p < 0.01). CONCLUSIONS: We here demonstrate that lateralized, remapped haptic feedback modulates motor awareness in a systematic, gait-cycle dependent manner. Based on this approach, the FeetBack system was used to provide augmented sensory information pertinent to the user's on-going movement such that they reported high motor awareness for (re)synchronized feedback of their movements. While motor adaptation was limited in the current cohort of healthy participants, the next step will be to evaluate if individuals with a compromised peripheral nervous system, as well as those with conditions of the central nervous system such as Parkinson's Disease, may benefit from the FeetBack system, both for maintaining a sense of agency over their movements as well as for systematic gait-adaptation in response to the remapped, self-paced, rhythmic feedback.

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The sense of agency is the ability to recognize that we are the actors of our actions and their consequences. We explored whether and how spatial cues may modulate the agency experience by manipulating the ecological validity of the experimental setup (real-space or computer-based setup) and the distance of the action-outcome (near or far). We tested 58 healthy adults collecting explicit agency judgments and the perceived time interval between movements and outcomes (to quantify the intentional binding phenomenon, an implicit index of agency). Participants show greater implicit agency for voluntary actions when there is a temporal and spatial action-outcome contingency. Conversely, participants reported similar explicit agency for outcomes appearing in the near and far space. Notably, these effects were independent of the ecological validity of the setting. These results suggest that spatial proximity, realistic or illusory, is essential for feeling implicitly responsible for the consequences of our actions.

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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.

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Motor awareness is a complex, multifaceted construct involving the awareness of both (i) one's motor state while executing a movement or remaining still and (ii) one's motor abilities. The analysis of neurological syndromes associated with motor disorders suggests the existence of various different components which are, however, integrated into a model of motor awareness. These components are: (i) motor intention, namely, a conscious desire to perform an action; (ii) motor monitoring and error recognition, that is, the capacity to check the execution of the action and identify motor errors; and (iii) a general awareness of one's own motor abilities and deficits, that is, the capacity to recognize the general state of one's motor abilities about the performance of specific actions and the potential consequences of motor impairment. Neuroanatomical correlates involving the parietal and insular cortices, the medial and lateral frontal regions, and subcortical structures (basal ganglia and limbic system) support this multi-component model. Specific damage (or disconnections) to these structures results in a number of different disorders in motor awareness, such as anosognosia for hemiplegia and apraxia, and a number of symptoms which are specific to motor intention disorders (e.g., the Anarchic Hand Syndrome and Tourette's Syndrome) or motor monitoring (e.g., Parkinson's and Huntington's diseases). All of these clinical conditions are discussed in the light of a motor awareness model.

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INTRODUCTION: In patients with Parkinson's disease (PD), impulsivity is still a matter of investigation. It has been hypothesized that impulsive personality traits may favour impulse control disorder (ICD) onset during dopaminergic therapy. In healthy subjects, a relationship between the awareness of motor intention and impulsive personality traits assessed by the Barratt impulsivity scale (BIS-11) has been reported. The aim of this study was to evaluate the relationship between the awareness of voluntary action and impulsivity traits in PD. METHODS: Twenty-eight PD patients (stages I-III on the Hoehn and Yahr scale) underwent an impulsivity trait assessment by the BIS-11 scale and a task based on the Libet's clock. Participants were requested to perform a self-initiated movement and report the time they first feel their intention to move (W-judgement) or the time of the actual movement (M-judgement). RESULTS: In patients with higher BIS-11 scores, the time lag between the W-judgement and the actual movement was significantly lower than in patients with lower BIS-11. No difference emerged in the M-judgement. CONCLUSION: Data suggest that also in PD patients, the impulsive personality trait is related to a "delayed" awareness of motor intention and therefore to a shorter interval to allow a conscious "veto" of the impending action. Characterization of the temporal profile of awareness of motor intention could prove useful in identifying PD patients at risk of developing ICDs during dopaminergic treatment.

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While outcomes of embodied awareness practices in terms of improved posture and flexibility, movement efficiency, and well-being are often reported, systematic investigations of such training effects and of the actual nature, extent, and neurofunctional correlates of learning mechanisms thought to lie at the core of such practices are very limited. The present study focused on the Feldenkrais method (FM), one of the most established embodied awareness practices, and aimed at investigating the neurofunctional outcomes of the somatic learning process at the core of the method by testing the modulations induced by a standardized FM protocol on the complexity of practicers' body structural map and on the activity of their sensorimotor network during different movement-related tasks (i.e., gestures observation, execution, and imagery). Twenty-five participants were randomly divided into an experimental group-which completed a 28-session FM protocol based on guided group practice-and a control group, and underwent pre-/post-training psychometric and electrophysiological assessment. Data analysis highlighted, at the end of the FM protocol, a significant increase of EEG markers of cortical activation (task-related mu desynchronization) in precentral regions during action observation and in central regions during action execution and imagery. Also, posterior regions of the sensorimotor network showed systematic activation during all the action-related tasks.

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The syndrome of anosognosia for hemiplegia, or the lack of awareness for one's paralysis following right hemisphere stroke, can provide unique insights into the neurocognitive mechanisms of self-awareness. Yet it remains unclear whether anosognosia for hemiplegia is a modality-specific deficit of sensorimotor monitoring, or whether domain-general processes of attention and belief-updating converge to cause anosognosia for hemiplegia. Using a Bayesian learning framework, we formalized and empirically investigated the hypothesis that failures to update anosognosic beliefs can be explained by abnormalities in the relative uncertainty (i.e. precision) ascribed to prior beliefs versus sensory information in different contexts. We designed a new motor belief-updating task that manipulated both the temporal (prospective and retrospective) and spatial (hemispace most affected by inattention and hemispace less affected by inattention) conditions in which beliefs had to be updated, and we validated its sensitivity to anosognosia for hemiplegia in 26 patients with right hemisphere stroke. We then computed and empirically tested two different Bayesian predictors of prospective beliefs using two proxies for precision in anosognosia for hemiplegia patients: (i) standardized, neuropsychological measures of objective attention abilities, i.e. visuospatial neglect scores and (ii) subjective uncertainty reports, i.e. confidence ratings. Our results suggest that while neglect does not affect local, sensorimotor error monitoring, it does seem to affect the degree to which observed errors are used to update more general, prospective beliefs about counterfactual motor abilities in anosognosia for hemiplegia. Difficulties in such 'counterfactual' belief-updating were associated with disruptions in tracts of the ventral attentional network (i.e. superior longitudinal fasciculus connecting the temporo-parietal junction and ventral frontal cortex) and associated lesions to the insula, inferior parietal cortex and superior temporal regions. These results suggest that self-awareness extends beyond local, retrospective monitoring, requiring also salience-based, convergence of beliefs about the self that go beyond the 'here-and-now' of sensorimotor experience.

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Introduction: Anosognosia for hemiplegia (AHP) is a condition in which patients with paralysis are unaware of their motor deficits. Research into AHP is important for improving its treatment and providing insight into the neurocognitive mechanism of motor awareness. Unfortunately, most studies use assessments with widely recognized limitations.The study aims at developing a psychometrically validated assessment of AHP.Method: We developed a 40-item Motor Unawareness Assessment (MUNA) and administered it to 131 right-hemisphere stroke patients. Principal Component Analysis (PCA) was used to identify the underlying factor structure. Receiver Operating Characteristics (ROC) analysis was used to determine diagnostic cutoffs, and Area Under the Curve (AUC) analysis used to assess these cutoffs. Relationships with demographic, clinical and neuropsychological variables were explored.Results: Five factors were identified: explicit motor awareness, implicit motor awareness, impaired sense of ownership, agency and illusory movement, and emotional reactions. Established cutoffs had excellent sensitivity and specificity. Clinical, neuropsychological and demographic variables did not predict overall MUNA score but were related to specific subcomponents.Conclusion: The MUNA can differentiate various facets of AHP and provides a detailed profile of (un)awareness. The MUNA can therefore provide robust assessment for research purposes and assist clinicians when developing targeted rehabilitation.

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Sense of agency refers to the feeling that one's self-generated action caused an external environment event. In a previous study, we suggested that the supplementary motor area (SMA), in its anterior portion (pre-SMA), is a key structure for attributing the sense of agency for the visual consequences of self-generated movements. However, real-life actions can lead to outcomes in different sensory modalities, raising the question of whether SMA represents a supra-modal hub for the sense of agency. Here, we compared the agency experience for visual and auditory outcomes by taking advantage of the intentional binding effect (IB). We observed discrete time-windows for the agency manifestation across different sensory modalities: While there was an IB at 200 ms delay between the action and the visual outcome, a time compression was observed when the auditory outcome followed the action by 400 ms. The magnitude of the IB was mirrored by meaningful brain activity in the pre-SMA but only at the specific delay when a sizeable IB was seen. We conclude that attributing consequences of self-generated movements to one's action is based on similar mechanisms across sensory modalities and that those mechanisms are related to the functioning of the motor system.

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Electrophysiological and neuroimaging studies suggest that our actions are initiated by unconscious mental processes long before awareness of intention to act. The time window between the awareness of the intention to move and the movement onset, which normally permits to exert a conscious "veto" on the impending action, is modulated by individual differences in trait impulsivity. In particular, trait impulsive people show a delayed awareness of the intention to act, probably exceeding the "point of no return", after which the action can no longer be inhibited. In order to investigate if individual differences in the "veto" interval might be explained by differences in the readiness potential (RP) dynamics, nineteen healthy participants underwent an impulsivity trait assessment using the Barratt Impulsiveness Scale (BIS-11) and performed a task based on Libet's clock paradigm, during EEG recordings of pre-movement neural activity. We observed a positive relationship between impulsive personality trait and motor system excitability during the preparation of self-initiated movements. In particular, the RP showed an earlier negative rising phase and a greater amplitude, with the increasing of BIS-11 scores. Based on present results, we conclude hypothesizing that trait impulsivity might be characterized by less effective preparatory inhibition mechanisms, which have a fundamental role in the control of behavior.

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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.

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The various neurocognitive processes contributing to the sense of body ownership have been investigated extensively in healthy participants, but studies in neurological patients can shed unique light into such phenomena. Here, we aimed to investigate whether visual capture by a fake hand (without any synchronous or asynchronous tactile stimulation) affects body ownership in a group of hemiplegic patients with or without disturbed sensation of limb ownership (DSO) following damage to the right hemisphere. We recruited 31 consecutive patients, including seven patients with DSO. The majority of our patients (64.5% overall and up to 86% of the patients with DSO) experienced strong feelings of ownership over a rubber hand within 15 sec following mere visual exposure, which correlated with the degree of proprioceptive deficits across groups and in the DSO group. Using voxel-based lesion-symptom mapping analysis, we were able to identify lesions associated with this pathological visual capture effect in a selective fronto-parietal network, including significant voxels (p < .05) in the frontal operculum and the inferior frontal gyrus. By contrast, lesions associated with DSO involved more posterior lesions, including the right temporoparietal junction and a large area of the supramarginal gyrus, and to a lesser degree the middle frontal gyrus. Thus, this study suggests that our sense of ownership includes dissociable mechanisms of multisensory integration.

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Anosognosia for hemiplegia (AHP) is informative about the neurocognitive basis of motor awareness. However, it is frequently associated with concomitant symptoms, such as hemispatial neglect and disturbances in the sense of body ownership (DSO). Although double dissociations between these symptoms have been reported, there is ongoing debate about whether they are manifestations of independent abnormalities, or a single neurocognitive deficit. We aimed to investigate the specificity of lesions associated with AHP by surpassing four, existing methodological limitations: (a) recruit a relatively large sample of patients (total N = 70) in a multi-centre study; (b) identify lesions associated with AHP in grey and white matter using voxel-based methods; (c) take into account the duration of AHP and concomitant neglect symptoms; and (d) compare lesions against a control hemiplegic group, patients suffering from AHP and DSO, and a few, rare patients with selective DSO. Results indicated that acute AHP is associated with a wide network, mainly including: (1) the Rolandic operculum, (2) the insula and (3) the superior temporal gyri. Subcortically, damage mainly involved the basal ganglia and white matter, mostly the superior corona radiate, arcuate fasciculus and the part of the ventral, superior longitudinal fasciculus. Persistent symptoms were linked with wider damage involving fronto-temporal cortex and long white matter tracts. A shift in the latero-medial direction (mainly involving the basal ganglia and surrounding white matter) emerged when DSO was taken accounted for. These results suggest that while bodily awareness is processed by areas widely distributed across the brain, intact subcortical structures and white matter tracts may be necessary to support basic feelings of owning and controlling contralateral body parts. An accurate and 'up-to-date' awareness of our motor abilities, however, may rely also on intact processing in cortical areas which presumably allow higher-order inferences about the current state of the body.

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Deficient voluntary control of behaviour and impulsivity are key aspects of impulse control disorders. The objective of the present study was to evaluate the relationship between behavioural measures of impulsivity and the awareness of voluntary action. Seventy-four healthy volunteers completed the Barratt Impulsiveness Scale (BIS), a questionnaire used to measure impulsive personality traits, and a go/no-go task. Moreover, all participants performed a behavioural task based on the Libet's clock paradigm in which they were requested to report the time of a self-initiated movement (M-judgement) or the time they first feel their intention to move (W-judgement). A positive relationship between the time in which subjects reported the intention to move (W-judgement) and impulsivity measures emerged. Namely, the higher was the score in the attentional and motor impulsivity subscales of the BIS and the number of inhibitory failure responses in the go/no-go task, the lower was the difference between the W-judgement and the actual movement (i.e. the awareness of intention to move was closer to the voluntary movement execution). In contrast, no relationship emerged with M-judgement. The present findings suggest that impulsivity is related to a delayed awareness of voluntary action. We hypothesize that in impulse control disorders, the short interval between conscious intention and actual movement may interfere with processes underlying the conscious 'veto' of the impending action.

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Here we investigated the temporal perception of self- and other-generated actions during sequential joint actions. Participants judged the perceived time of two events, the first triggered by the participant and the second by another agent, during a cooperative or competitive interaction, or by an unspecified mechanical cause. Results showed that participants perceived self-generated events as shifted earlier in time (anticipation temporal judgment bias) and non-self-generated events as shifted later in time (repulsion temporal judgment bias). This latter effect was observed independently from the kind of cause (i.e., agentive or mechanical) or interaction (i.e., cooperative or competitive). We suggest that this might represent a mental process which allows discriminating events that cannot plausibly be linked to one's own action. When an event immediately follows a self-generated one, temporal judgment biases operate as self-serving biases in order to separate self-generated events from events of another physical causality.

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In their seminal (1983) study, Libet and colleagues suggested that awareness of one's intention to act has a postdictive character in that it occurs long after cerebral activity leading to action has been initiated. Crucially, Libet et al. further suggested that the time window (±200 ms) between the conscious experience of the intention to act and the action itself offers people the possibility of "vetoing" the unfolding action. This raises the question of whether there are individual differences in the duration of this "veto window" and which components of the readiness potential (RP) and the lateralized readiness potential (LRP) explain this variability. It has been reported that some psychiatric diseases lead to shorter intervals between conscious intentions and actions. However, it is unclear whether such patients suffer from impairment of the sense of volition, thus experiencing voluntary movements as involuntary, or whether voluntary inhibition of action is actually reduced, since conscious intention occurs later. We had two aims in the present paper. First, we aimed at clarifying the role of consciousness in voluntary actions by examining the relation between the duration of the veto window and impulsivity. Second, we sought to examine different components of the RP and LRP waveforms so as to attempt to explain observed variability in W judgments. Our results indicate (1) that impulsive people exhibit a shorter delay between their intention and the action than non-impulsive people, and (2) that this difference can hardly be attributed to a difference in time perception. Electroencephalography indicated that the rate of growth of the RP is relevant to explain differences in W judgments, since we observed that the RP at the moment of conscious intention is lower for people with late conscious intention than for people with early conscious intention. The onset and the intercept of these waveforms were less interpretable. These results bring new light on the role that consciousness plays in voluntary action.

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Anarchic hand syndrome (AHS) is characterized by goal-directed movements performed without volitional control (agency). Different AHS subtypes have been identified; however, few studies have examined the posterior subtype. We report a case of AHS following right-hemisphere parietal damage, with left-sided somatosensory and proprioceptive impairment. Agency was examined for nonanarchic (volitional) movements performed using the anarchic hand. The patient experienced abnormal agency for movements whether motor intention and visual feedback were congruent or incongruent, but not when intention was absent (passive movement). Findings suggest a general disturbance of veridical motor awareness and agency in this case of parietal AHS.

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Anosognosia for hemiplegia (AHP), or unawareness of motor deficits contralateral to a brain lesion, has lasting negative implications for the management and rehabilitation of patients. A recent, bedside psychophysical intervention, namely self-observation by video replay, lead to a lasting remission of severe AHP in an acute stroke patient (Fotopoulou, A., Rudd, A., Holmes, P., & Kopelman, M. (2009). Self-observation reinstates motor awareness in anosognosia for hemiplegia. Neuropsychologia, 47, 1256-1260). This procedure has been adjusted and applied here, as the basis of two intervention protocols administered independently to two patients with severe AHP. The first study used multiple, successive sessions of video-based self-observation in an acute patient, targeting first the awareness of upper limb and subsequently lower limb paralysis. The second study used a single session of video-based, self- and other-observation in a patient at the chronic stage following onset. Both protocols also involved elements of rapport building and emotional support. The results revealed that video-based self-observation had dramatic, immediate effects on awareness in both acute and chronic stages and it seemed to act as an initial trigger for eventual symptom remission. Nevertheless, these effects did not automatically generalise to all functional domains. This study provides provisional support that video-based self-observation may be included in wider rehabilitation programmes for the management and restoration of anosognosia.

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Right brain damage patients may not complain of a left sided paralysis up to the point of denying it or even claiming of having just moved an otherwise paralyzed limb. This condition is known as anosognosia for hemiplegia (AHP). Recent behavioural experiments suggest that some residual intentionality might be preserved in patients with anosognosia and that the false belief of having moved originates from a failure to notice discrepancies between movement expectancies and the actual state of the motor system. This failure may be caused by a lack of afferent sensory information concerning the movement or alternatively by a direct dysfunction of the brain regions involved in actions' motor monitoring (i.e., the comparator system). Here we examined the effect of anodal transcranial direct current stimulation (tDCS) of the right premotor cortex in a patient with a bilateral lesion, involving predominantly the right hemisphere, and a dense unawareness for his left hemiplegia. During sham or anodal tDCS the patient was requested to judge his ability to perform simple motor actions (i) without actually executing the movement itself ("offline" condition) and after having performed a series of verbally cued finger opposition movements ("online" condition) with (i) eyes-closed or (ii) eyes-open. We found that anodal tDCS induces a significant remission of the false experience of movement only when the patient is requested to actually perform the movement with eyes open. Conversely, the patient's awareness does not improve in both the "offline" condition (in which the patient does not attempt to perform the movement) and in the "online" condition, when vision is precluded ("online" condition, eyes-closed). We conclude that the stimulation of the premotor cortex by tDCS activates brain regions involved in motor monitoring, temporary restoring the ability of the motor comparator system to correctly appreciate afferent information and build up a veridical motor awareness.

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The possible role of emotion in anosognosia for hemiplegia (i.e., denial of motor deficits contralateral to a brain lesion), has long been debated between psychodynamic and neurocognitive theories. However, there are only a handful of case studies focussing on this topic, and the precise role of emotion in anosognosia for hemiplegia requires empirical investigation. In the present study, we aimed to investigate how negative and positive emotions influence motor awareness in anosognosia. Positive and negative emotions were induced under carefully-controlled experimental conditions in right-hemisphere stroke patients with anosognosia for hemiplegia (n = 11) and controls with clinically normal awareness (n = 10). Only the negative, emotion induction condition resulted in a significant improvement of motor awareness in anosognosic patients compared to controls; the positive emotion induction did not. Using lesion overlay and voxel-based lesion-symptom mapping approaches, we also investigated the brain lesions associated with the diagnosis of anosognosia, as well as with performance on the experimental task. Anatomical areas that are commonly damaged in AHP included the right-hemisphere motor and sensory cortices, the inferior frontal cortex, and the insula. Additionally, the insula, putamen and anterior periventricular white matter were associated with less awareness change following the negative emotion induction. This study suggests that motor unawareness and the observed lack of negative emotions about one's disabilities cannot be adequately explained by either purely motivational or neurocognitive accounts. Instead, we propose an integrative account in which insular and striatal lesions result in weak interoceptive and motivational signals. These deficits lead to faulty inferences about the self, involving a difficulty to personalise new sensorimotor information, and an abnormal adherence to premorbid beliefs about the body.

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