Multisensory processing is enhanced for stimuli that appear in close proximity to our body. The so-called peripersonal space (PPS) not only serves as an evolutionary, protective mechanism but also localizes and demarcates our self in space. While our perceived self-location is generally unitary and firmly located at the position of our body, there are rare clinical instances in which patients report the presence of a double (heautoscopy), an actual duplication of the self, and experience themselves to be located simultaneously both in their physical body and in that of the double. Here we used augmented reality and drone technology to create a heautoscopic view of one’s own double: Participants were presented with real-time feedback from a drone-mounted camera that autonomously followed them as they responded to multisensory stimuli on their body and in their environment. Results demonstrate that such manipulation induced a heautoscopy-like state in healthy participants, characterized by two co-existing but spatially distinct PPSs: tactile processing on the body was enhanced by visual stimuli approaching both the participant’s body and, in case of on-going locomotor navigation, the location of their double. While previous studies have shown that PPS remaps towards the location of actions or the perceived self-location, this is the first evidence that PPS can co-exist in two locations.
Hallucinations can occur in the healthy population, are clinically relevant and frequent symptoms in many neuropsychiatric conditions, and have been shown to mark disease progression in patients with neurodegenerative disorders where antipsychotic treatment remains challenging. Here, we combine MR-robotics capable of inducing a clinically-relevant hallucination, with real-time fMRI neurofeedback (fMRI-NF) to train healthy individuals to up-regulate a fronto-parietal brain network associated with the robotically-induced hallucination. Over three days, participants learned to modulate occurrences of and transition probabilities to this network, leading to heightened sensitivity to induced hallucinations after training. Moreover, participants who became sensitive and succeeded in fMRI-NF training, showed sustained and specific neural changes after training, characterized by increased hallucination network occurrences during induction and decreased hallucination network occurrences during a matched control condition. These data demonstrate that fMRI-NF modulates specific hallucination network dynamics and highlights the potential of fMRI-NF as a novel antipsychotic treatment in neurodegenerative disorders and schizophrenia. fMRI-neurofeedback targeting a hallucination-related whole-brain network allows individuals to willfully control its temporal properties and modify proneness the hallucination induction, showing promise for investigating future therapies in disease.
The use of hands for gathering rich sensory information is essential for proper interaction with the environment; therefore, the restoration of sensation is critical for reestablishing the sense of embodiment in hand amputees. Here, we show that a noninvasive wearable device can be used to provide thermal sensations on amputees' phantom hands. The device delivers thermal stimuli to specific regions of skin on their residual limb. These sensations were phenomenologically similar to those on the intact limbs and were stable over time. Using the device, the subjects could successfully exploit the thermal phantom hand maps to detect and discriminate different thermal stimuli. The use of a wearable device that provides thermal sensation can increase the sense of embodiment and improve life quality in hand amputees.
Visuo-motor integration shapes our daily experience and underpins the sense of feeling in control over our actions. The last decade has seen a surge in robotically and virtually mediated interactions, whereby bodily actions ultimately result in an artificial movement. But despite the growing number of applications, the neurophysiological correlates of visuo-motor processing during human-machine interactions under dynamic conditions remain scarce. Here we address this issue by employing a bimanual robotic interface able to track voluntary hands movement, rendered in real-time into the motion of two virtual hands. We experimentally manipulated the visual feedback in the virtual reality with spatial and temporal conflicts and investigated their impact on (1) visuo-motor integration and (2) the subjective experience of being the author of one's action (i.e., sense of agency). Using somatosensory evoked responses measured with electroencephalography, we investigated neural differences occurring when the integration between motor commands and visual feedback is disrupted. Our results show that the right posterior parietal cortex encodes for differences between congruent and spatially-incongruent interactions. The experimental manipulations also induced a decrease in the sense of agency over the robotically-mediated actions. These findings offer solid neurophysiological grounds that can be used in the future to monitor integration mechanisms during movements and ultimately enhance subjective experience during human-machine interactions.
Although hallucinations are important and frequent symptoms in major psychiatric and neurological diseases, little is known about their brain mechanisms. Hallucinations are unpredictable and private experiences, making their investigation, quantification and assessment highly challenging. A major shortcoming in hallucination research is the absence of methods able to induce specific and short-lasting hallucinations, which resemble clinical hallucinations, can be elicited repeatedly and vary across experimental conditions. By integrating clinical observations and recent advances in cognitive neuroscience with robotics, we have designed a novel device and sensorimotor method able to repeatedly induce a specific, clinically relevant hallucination: presence hallucination. Presence hallucinations are induced by applying specific conflicting (spatiotemporal) sensorimotor stimulation including an upper extremity and the torso of the participant. Another, MRI-compatible, robotic device using similar sensorimotor stimulation permitted the identification of the brain mechanisms of these hallucinations. Enabling the identification of behavioral and a frontotemporal neural biomarkers of hallucinations, under fully controlled experimental conditions and in real-time, this method can be applied in healthy participants as well as patients with schizophrenia, neurodegenerative disease or other hallucinations. The execution of these protocols requires intermediate-level skills in cognitive neuroscience and MRI processing, as well as minimal coding experience to control the robotic device. These protocols take ~3 h to be completed.
The perception that someone is nearby, although nobody can be seen or heard, is called presence hallucination (PH). Being a frequent hallucination in patients with Parkinson's disease, it has been argued to be indicative of a more severe and rapidly advancing form of the disease, associated with psychosis and cognitive decline. PH may also occur in healthy individuals and has recently been experimentally induced, in a controlled manner during fMRI, using MR-compatible robotics and sensorimotor stimulation. Previous neuroimaging correlates of such robot-induced PH, based on conventional time-averaged fMRI analysis, identified altered activity in the posterior superior temporal sulcus and inferior frontal gyrus in healthy individuals. However, no link with the strength of the robot-induced PH was observed, and such activations were also associated with other sensations induced by robotic stimulation. Here we leverage recent advances in dynamic functional connectivity, which have been applied to different psychiatric conditions, to decompose fMRI data during PH-induction into a set of co-activation patterns that are tracked over time, as to characterize their occupancies, durations, and transitions. Our results reveal that, when PH is induced, the identified brain patterns significantly and selectively increase their transition probabilities towards a specific brain pattern, centred on the posterior superior temporal sulcus, angular gyrus, dorso-lateral prefrontal cortex, and middle prefrontal cortex. This change is not observed in any other control conditions, nor is it observed in association with other sensations induced by robotic stimulation. The present findings describe the neural mechanisms of PH in healthy individuals and identify a specific disruption of the dynamics of network interactions, extending previously reported network dysfunctions in psychotic patients with hallucinations to an induced robot-controlled specific hallucination in healthy individuals.
A growing number of studies have focused on identifying cognitive processes that are modulated by interoceptive signals. Here we investigated whether interoception affects self-processing, by assessing changes in self-voice perception as a function of respiratory and cardiac cycles. Considering the fundamental role interoception plays in bodily self-consciousness, we additionally applied conflicting sensorimotor stimulation inducing a state characterized by a loss of self and increased otherness, and investigated its effects in self-other voice perception. Our data reveal that breathing, but not heartbeat, affects self-voice perception, by showing that participants (N = 30) discriminated self-voice from other voices better during inspiration, while being in the state of increased otherness and especially when hearing voices of other people. Loudness judgement of equivalent self-related stimuli was unaffected by breathing. Combining interoception and voice perception with self-monitoring framework, these data extend recent findings on breathing-dependent cognition to self-processing.
Psychosis, characterized by hallucinations and delusions, is a common feature of psychiatric disease, especially schizophrenia. One prominent theory posits that psychosis is driven by abnormal sensorimotor predictions leading to the misattribution of self-related events. This misattribution has been linked to passivity experiences (PE), such as loss of agency and, more recently, to presence hallucinations (PH), defined as the conscious experience of the presence of an alien agent while no person is actually present. PH has been observed in schizophrenia, Parkinson's disease, and neurological patients with brain lesions and, recently, the brain mechanisms of PH (PH-network) have been determined comprising bilateral posterior middle temporal gyrus (pMTG), inferior frontal gyrus (IFG), and ventral premotor cortex (vPMC). Given that the experience of an alien agent is a common feature of PE, we here analyzed the functional connectivity within the PH-network in psychotic patients with (N = 39) vs without PE (N = 26). We observed reduced fronto-temporal functional connectivity in patients with PE compared to patients without PE between the right pMTG and the right and left IFG of the PH-network. Moreover, when seeding from these altered regions, we observed specific alterations with brain regions commonly linked to auditory-verbal hallucinations (such as Heschl's gyrus). The present connectivity findings within the PH-network extend the disconnection hypothesis for hallucinations to the specific case of PH and associates the PH-network with key brain regions for frequent psychotic symptoms such as auditory-verbal hallucinations, showing that PH are relevant to the study of the brain mechanisms of psychosis and PE.
Previous studies have shown that self-generated stimuli in auditory, visual, and somatosensory domains are attenuated, producing decreased behavioral and neural responses compared with the same stimuli that are externally generated. Yet, whether such attenuation also occurs for higher-level cognitive functions beyond sensorimotor processing remains unknown. In this study, we assessed whether cognitive functions such as numerosity estimations are subject to attenuation in 56 healthy participants (32 women). We designed a task allowing the controlled comparison of numerosity estimations for self-generated (active condition) and externally generated (passive condition) words. Our behavioral results showed a larger underestimation of self-generated compared with externally generated words, suggesting that numerosity estimations for self-generated words are attenuated. Moreover, the linear relationship between the reported and actual number of words was stronger for self-generated words, although the ability to track errors about numerosity estimations was similar across conditions. Neuroimaging results revealed that numerosity underestimation involved increased functional connectivity between the right intraparietal sulcus and an extended network (bilateral supplementary motor area, left inferior parietal lobule, and left superior temporal gyrus) when estimating the number of self-generated versus externally generated words. We interpret our results in light of two models of attenuation and discuss their perceptual versus cognitive origins.SIGNIFICANCE STATEMENTWe perceive sensory events as less intense when they are self-generated compared with when they are externally generated. This phenomenon, called attenuation, enables us to distinguish sensory events from self and external origins. Here, we designed a novel fMRI paradigm to assess whether cognitive processes such as numerosity estimations are also subject to attenuation. When asking participants to estimate the number of words they had generated or passively heard, we found bigger underestimation in the former case, providing behavioral evidence of attenuation. Attenuation was associated with increased functional connectivity of the intraparietal sulcus, a region involved in numerosity processing. Together, our results indicate that the attenuation of self-generated stimuli is not limited to sensory consequences but is also impact cognitive processes such as numerosity estimations.
Hallucinations in Parkinson's disease (PD) are disturbing and frequent non-motor symptoms and constitute a major risk factor for psychosis and dementia. We report a robotics-based approach applying conflicting sensorimotor stimulation, enabling the induction of presence hallucinations (PHs) and the characterization of a subgroup of patients with PD with enhanced sensitivity for conflicting sensorimotor stimulation and robot-induced PH. We next identify the fronto-temporal network of PH by combining MR-compatible robotics (and sensorimotor stimulation in healthy participants) and lesion network mapping (neurological patients without PD). This PH-network was selectively disrupted in an additional and independent cohort of patients with PD, predicted the presence of symptomatic PH, and associated with cognitive decline. These robotics-neuroimaging findings extend existing sensorimotor hallucination models to PD and reveal the pathological cortical sensorimotor processes of PH in PD, potentially indicating a more severe form of PD that has been associated with psychosis and cognitive decline.
Spinal cord stimulation (SCS) is an approved treatment for truncal and limb neuropathic pain. However, pain relief is often suboptimal and SCS efficacy may reduce over time, sometimes requiring addition of other pain therapies, stimulator revision, or even explantation. We designed and tested a new procedure by combining SCS with immersive virtual reality (VR) to enable analgesia in patients with chronic leg pain. We coupled SCS and VR by linking SCS-induced paresthesia with personalized visual bodily feedback that was provided by VR and matched to the spatio-temporal patterns of SCS-induced paresthesia. In this cross-sectional prospective interventional study, fifteen patients with severe chronic pain and an SCS implant underwent congruent SCS-VR (personalized visual feedback of the perceived SCS-induced paresthesia displayed on the patient's virtual body) and two control conditions (incongruent SCS-VR; VR alone). We demonstrate the efficacy of neuromodulation-enhanced VR for the treatment of chronic pain by showing that congruent SCS-VR reduced pain ratings on average by 44%. SCS-VR analgesia was stronger than in both control conditions (enabling stronger analgesic effects than incongruent SCS-VR analgesia or VR alone), and kept increasing over successive stimulations, revealing the selectivity and consistency of the observed effects. We also show that analgesia persists after congruent SCS-VR had stopped, indicating carry over effects and underlining its therapeutic potential. Linking latest VR technology with recent insights from the neuroscience of body perception and SCS-neuromodulation, our personalized new SCS-VR platform highlights the impact of immersive digiceutical therapies for chronic pain.
Sensorimotor conflicts are known to alter the perception of accompanying sensory signals, and deficits in sensory attenuation have been observed in schizophrenia. In the auditory domain, self-generated tones or voices (compared to tones or voices presented passively or with temporal delays) have been associated with changes in loudness perception and attenuated neural responses. It has been argued that for sensory signals to be attenuated, predicted and sensory consequences must have a consistent spatiotemporal relationship, between button presses and reafferent signals, via predictive sensory signaling, a process altered in schizophrenia. Here, we investigated auditory sensory attenuation for a series of morphed voices while healthy participants applied sensorimotor stimulations that had no spatiotemporal relationship to the voice stimuli and that have been shown to induce mild psychosis-like phenomena. In two independent groups of participants, we report a loudening of silent voices and found this effect only during maximal sensorimotor conflicts (versus several control conditions). Importantly, conflicting sensorimotor stimulation also induced a mild psychosis-like state in the form of somatic passivity and participants who experienced stronger passivity lacked the sensorimotor loudening effect. We argue that this conflict-related sensorimotor loudness amplification may represent a reduction of auditory self-attenuation that is lacking in participants experiencing a concomitant mild psychosis-like state. We interpret our results within the framework of the comparator model of sensorimotor control, and discuss the implications of our findings regarding passivity experiences and hallucinations in schizophrenia.
Abstract Background The ability to recognize whether sensory consequences have been self-generated or externally produced is an important element of motor control and self-monitoring. Deficits in self-monitoring have been proposed to cause abnormal bodily experiences and psychotic symptoms such as hallucinations. A recent study designed a robotic system that applies sensorimotor stimulation in healthy subjects and safely induces mild presence hallucinations (PH) and passivity experiences (Blanke et al., 2014). PH are defined as the sensation that someone is close by when actually no one is present and passivity experiences are characterized by perceptions or beliefs that an external agent is controlling one’s actions, perceptions, and/or thoughts. Although, both symptoms occur in schizophrenia and Parkinson’s disease, their neural mechanisms are unknown. Methods Here, we first investigated the neural mechanisms of PH and passivity experiences in 25 healthy subjects. We developed a new MR-compatible robotic system able to generate the aforementioned sensorimotor conflicts, while recording subjects’ brain activity using fMRI. In addition, we applied lesion network mapping to 11 neurological patients with symptomatic PH and compared both populations to find a common PH network. In a final step, we investigated the relevance of the PH network by analyzing the connectivity of the PH network in resting state fMRI data from 58 psychotic patients. Results We first evaluated the regions associated with the general sensorimotor conflict, which revealed the left sensorimotor area, the left putamen, the right inferior parietal lobule and the right cerebellum. Then, we analyzed the regions that were more activated during the condition eliciting PH and passivity experiences and found the inferior frontal gyrus (IFG), the insula, the superior medial gyrus and the middle temporal gyrus (MTG). Comparison of these two networks with the symptomatic PH network in neurological patients highlighted the IFG, the MTG and the vPMc. The resting state analysis within those regions in psychotic patients revealed no global differences between the groups but a functional connectivity decrease between MTG and IFG (bilaterally) specific for psychotic patients experiencing passivity experiences. Discussion Collectively, we showed that through the use of a robotic system generating sensorimotor conflicts, the neural correlates of induced-PH and passivity experiences can be studied in healthy subjects in a controlled manner. In addition, we found two networks associated with induced-PH and passivity experience. Of these regions, three were also recruited in patients with PH of neurological origin, forming the PH network. MTG-IFG connectivity in the PH network was altered selectively in psychotic patients with passivity experiences, revealing the relevance of the neural mechanisms of PH and passivity experiences in psychosis.
Thought insertion (TI) is characterized by the experience that certain thoughts, occurring in one's mind, are not one's own, but the thoughts of somebody else and suggestive of a psychotic disorder. We report a robotics-based method able to investigate the behavioral and subjective mechanisms of TI in healthy participants. We used a robotic device to alter body perception by providing online sensorimotor stimulation, while participants performed cognitive tasks implying source monitoring of mental states attributed to either oneself or another person. Across several experiments, conflicting sensorimotor stimulation reduced the distinction between self- and other-generated thoughts and was, moreover, associated with the experimentally generated feeling of being in the presence of an alien agent and subjective aspects of TI. Introducing a new robotics-based approach that enables the experimental study of the brain mechanisms of TI, these results link TI to predictable self-other shifts in source monitoring and specific sensorimotor processes.
Summary Hallucinations in Parkinson’s disease (PD) are one of the most disturbing non-motor symptoms, affect half of the patients, and constitute a major risk factor for adverse clinical outcomes such as psychosis and dementia. Here we report a robotics-based approach, enabling the induction of a specific clinically-relevant hallucination (presence hallucination, PH) under controlled experimental conditions and the characterization of a PD subgroup with enhanced sensorimotor sensitivity for such robot-induced PH. Using MR-compatible robotics in healthy participants and lesion network mapping analysis in neurological non-PD patients, we identify a fronto-temporal network that was associated with PH. This common PH-network was selectively disrupted in a new and independent sample of PD patients and predicted the presence of symptomatic PH. These robotics-neuroimaging findings determine the behavioral and neural mechanisms of PH and reveal pathological cortical sensorimotor processes of PH in PD, identifying a more severe form of PD associated with psychosis and cognitive decline.
Psychosis is an abnormal mental state including hallucinations and delusions, typical of psychiatric conditions such as schizophrenia. Recent theories posit that psychosis is driven by inaccurate sensorimotor predictions causing the misattribution of self-related events to external sources. This misattribution has been linked to first-rank symptoms (FRS), including a loss of sense of agency and, particularly, the occurrence of an alien agent. The subjective experience of an alien agent while no one is actually there is called presence hallucination (PH). PH has been observed in schizophrenia, Parkinson’s disease and neurological patients with circumscribed brain lesions. In healthy participants, PH can be induced by exerting sensorimotor conflicts between the participants’ upper-limb movements and a tactile feedback received on the back using MRI-compatible robotics. Crucially, the neural network associated with this robotically-induced version of PH overlap with the symptomatic-PH network derived from neurological patients suffering from PH in right dorso-lateral prefrontal cortex (dlPFC) and middle temporal gyrus (MTG), suggesting a common neural mechanism (PH network). Given that experiencing an alien agent is a specific feature of FRS, we tested whether bilateral functional connectivity in PH network comprising of dlPFC and MTG can specifically differentiate psychotic patients with (N = 39) versus without (N = 24) FRS. We observed reduced functional connectivity in patients with FRS as compared to patients without FRS between the right MTG and the dlPFC bilaterally. Interestingly, connectivity between these areas was negatively correlated with the FRS severity (rho = -0.29 and rho = -0.26, respectively). We propose that reduced functional connectivity between the right MTG and bilateral dlPFC areas could be a specific biomarker of first rank symptoms in patients with psychosis.
Recent developments in virtual reality and robotic technologies have allowed investigating the behavioural and brain mechanisms that grounds self-consciousness in the multi-sensory (e.g. vision and touch) and sensorimotor processing of bodily signals. Yet, previous technological solutions to apply tactile stimuli for body illusion induction limit participants movements, do not allow for stimulations in dynamic environments (e.g., the subject walking), and can hardly be integrated into real-life settings and complex, interactive, virtual reality environments. Here, we present the development and first validation of a new semi-wearable haptic system, based on vibration technology, to induce a range of bodily illusions that are of relevance for research in psychiatry. This is a first step towards the development of wearable haptic systems able to administer touch and induce specific bodily illusions under dynamic conditions and in real-life settings.
Psychosis is often depicted as a disruption of the self-model. Patients suffering from psychosis report many symptoms relating to deficiencies in the minimal self, including loss of the sense of control over their actions (Sense of Agency) as well as numerous disturbances of body representation (e.g. Body Ownership). Positive symptoms of psychosis such as passivity symptoms and auditory hallucinations (termed first-rank symptoms) are characterized by a diminished demarcation of self-other boundaries, causing misattribution of self-generated actions to external sources. It has been suggested that this deficiency in self-monitoring in schizophrenia is due to abnormal sensorimotor prediction mechanisms, causing a loss of agency for actions and thoughts. While the neurobiological underpinnings of schizophrenia are yet unclear, many studies have reported aberrant neural connectivity in schizophrenia patients which may impact sensorimotor prediction and integration. Recently, we have shown that introducing sensorimotor conflict (SMC) can induce psychosis like symptoms in healthy patients. Employing a master-slave robotic system we induced a conflict by introducing a delay between the participants’ movements and the haptic feedback. The SMC caused a feeling of a Presence (FoP) which is a first rank symptom of psychosis. The FoP is also found in neurological patients with lesions in cortical regions of the temporoparietal cortex, insular cortex and fronto-parietal cortex related to abnormal bodily self-representation. Here, we tested if SMC may cause misattribution of auditory stimuli and if this is related to neural connectivity. We tested first episode psychosis patients (N=31) with and without first rank symptoms (related to the sense of control over actions and thoughts) as well as healthy participants (N=20) on an auditory attribution task while inducing conflict with the master-slave robot. We found that when a SMC was introduced patients with first rank symptoms showed a decrease in their ability to judge if the auditory stimuli were in their own voice or the voice of another person. Resting state functional connectivity analysis indicated that the first rank patients had reduced connectivity in the network related to the SMC, but not in other control regions. Furthermore, the reduced functional connectivity correlated with the rates of auditory misattribution. Our results show that induction of SMC can cause auditory misattributions, and that this is related to reduced cortical communication in regions related to sensorimotor body representation. These findings connect two influential theories of psychosis linking cognitive theories of sensorimotor prediction error in schizophrenia with systems level theories of neural disconnectivity. Understanding these neurocognitive mechanisms underlying the disruption of the self-model in psychosis may allow novel approaches in early diagnosis and treatment of these conditions.
Dysfunction of sensorimotor predictive processing is thought to underlie abnormalities in self-monitoring producing passivity symptoms in psychosis. Experimentally induced sensorimotor conflict can produce a failure in bodily self-monitoring (presence hallucination [PH]), yet it is unclear how this is related to auditory self-monitoring and psychosis symptoms. Here we show that the induction of sensorimotor conflict in early psychosis patients induces PH and impacts auditory-verbal self-monitoring. Participants manipulated a haptic robotic system inducing a bodily sensorimotor conflict. In experiment 1, the PH was measured. In experiment 2, an auditory-verbal self-monitoring task was performed during the conflict. Fifty-one participants (31 early psychosis patients, 20 matched controls) participated in the experiments. The PH was present in all participants. Psychosis patients with passivity experiences (PE+) had reduced accuracy in auditory-verbal self-other discrimination during sensorimotor stimulation, but only when sensorimotor stimulation involved a spatiotemporal conflict (F(2, 44) = 6.68, P = .002). These results show a strong link between robotically controlled alterations in sensorimotor processing and auditory misattribution in psychosis and provide evidence for the role of sensorimotor processes in altered self-monitoring in psychosis.
A major goal of neuroprosthetics is to design artificial limbs that are experienced (‘embodied’) like real limbs. However, despite important technological advances, this goal has not been reached and prosthesis embodiment is still very limited. Differently from our physical body, current bionic limbs do not provide the continuous multisensory feedback required for a limb to be experienced as one’s own. Here, we present a novel neuroprosthetic approach that combines peripheral neurotactile stimulation—inducing tactile sensation on the missing limb—and immersive digital technology—providing visual illumination of the prosthetic hand. We tested whether coherent multisensory visuo-tactile neural stimulation (VTNS)1 induced higher prosthesis embodiment and reduced the distorted perception of the phantom limb (telescoping, ie, the phantom limb is perceived as shorter than the intact limb). Patient 1 and patient 2 are transradial left forearm chronic amputees, who suffered upper limb telescoping. Patients were implanted with transverse intrafascicular multichannel electrodes (TIMEs), which induced the sensation of a vibration in a circumscribed skin region of the finger 2 via medial nerve stimulation in patient 1 (online supplementary figure 1A) and in a skin region of finger 5 via ulnar nerve stimulation in patient 2 (online supplementary figure 1B and material 1). Neurotactile stimulation2 was coupled with automatised visual illumination of a skin region on the patient’s prosthetic hand that corresponded to the somatotopic location of touch sensations experienced on the phantom hand (VTNS; online supplementary video 1, online supplementary figure 1, online supplementary material 1). VTNS was administered in two conditions, either with synchronous visual and neurotactile stimulation or in a control condition of asynchronous stimulation (1.5–2.5s delay). ### Supplementary data [jnnp-2018-318570-supp1.pdf] ### Supplementary video [jnnp-2018-318570-supp1.mp4] Prosthesis embodiment was measured via a questionnaire, whereas changes in phantom limb perception were tested via a body landmark task where patients indicated the perceived position of different parts of the phantom limb …