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1.  Touching a rubber hand: feeling of body ownership is associated with activity in multisensory brain areas 
In the ‘rubber hand illusion’, the sight of brushing of a rubber hand at the same time as brushing of the person’s own hidden hand is sufficient to produce a feeling of ownership of the fake hand. We have previously shown that this illusion is associated with activity in the multisensory areas, most notably the ventral premotor cortex (Ehrsson et al. Science 305:875–877, 2004). However, it remains to be demonstrated that this illusion does not simply reflect the dominant role of vision, and that the premotor activity does not reflect a visual representation of an object near the hand. To address these issues we introduce a somatic ‘rubber hand illusion’. The experimenter moved the blindfolded participants’ left index finger so that it touched the fake hand, and simultaneously, he touched the participants’ real right hand synchronizing the touches as perfectly as possible. After about 9.7 seconds this stimulation elicited an illusion that one was touching one’s own hand. We scanned brain activity during this illusion and two control conditions, using functional magnetic resonance imaging. Activity in the ventral premotor cortices, intraparietal cortices, and the cerebellum was associated with the illusion of touching one’s own hand. Furthermore, the rated strength of the illusion correlated with the degree of premotor and cerebellar activity. This finding suggests that the activity in these areas reflects the detection of congruent multisensory signals from one’s own body, rather than of visual representations. We propose that this could be the mechanism for the feeling of body ownership.
PMCID: PMC1395356  PMID: 16280594
premotor cortex; posterior parietal cortex; cerebellum; rubber hand illusion; functional magnetic resonance imaging; body image; self recognition
2.  The Importance of Synchrony and Temporal Order of Visual and Tactile Input for Illusory Limb Ownership Experiences – An fMRI Study Applying Virtual Reality 
PLoS ONE  2014;9(1):e87013.
In the so-called rubber hand illusion, synchronous visuotactile stimulation of a visible rubber hand together with one's own hidden hand elicits ownership experiences for the artificial limb. Recently, advanced virtual reality setups were developed to induce a virtual hand illusion (VHI). Here, we present functional imaging data from a sample of 25 healthy participants using a new device to induce the VHI in the environment of a magnetic resonance imaging (MRI) system. In order to evaluate the neuronal robustness of the illusion, we varied the degree of synchrony between visual and tactile events in five steps: in two conditions, the tactile stimulation was applied prior to visual stimulation (asynchrony of −300 ms or −600 ms), whereas in another two conditions, the tactile stimulation was applied after visual stimulation (asynchrony of +300 ms or +600 ms). In the fifth condition, tactile and visual stimulation was applied synchronously. On a subjective level, the VHI was successfully induced by synchronous visuotactile stimulation. Asynchronies between visual and tactile input of ±300 ms did not significantly diminish the vividness of illusion, whereas asynchronies of ±600 ms did. The temporal order of visual and tactile stimulation had no effect on VHI vividness. Conjunction analyses of functional MRI data across all conditions revealed significant activation in bilateral ventral premotor cortex (PMv). Further characteristic activation patterns included bilateral activity in the motion-sensitive medial superior temporal area as well as in the bilateral Rolandic operculum, suggesting their involvement in the processing of bodily awareness through the integration of visual and tactile events. A comparison of the VHI-inducing conditions with asynchronous control conditions of ±600 ms yielded significant PMv activity only contralateral to the stimulation site. These results underline the temporal limits of the induction of limb ownership related to multisensory body-related input.
PMCID: PMC3909015  PMID: 24498012
3.  Disturbances in Body Ownership in Schizophrenia: Evidence from the Rubber Hand Illusion and Case Study of a Spontaneous Out-of-Body Experience 
PLoS ONE  2011;6(10):e27089.
A weakened sense of self may contribute to psychotic experiences. Body ownership, one component of self-awareness, can be studied with the rubber hand illusion (RHI). Watching a rubber hand being stroked while one's unseen hand is stroked synchronously can lead to a sense of ownership over the rubber hand, a shift in perceived position of the real hand, and a limb-specific drop in stimulated hand temperature. We aimed to assess the RHI in schizophrenia using quantifiable measures: proprioceptive drift and stimulation-dependent changes in hand temperature.
The RHI was elicited in 24 schizophrenia patients and 21 matched controls by placing their unseen hand adjacent to a visible rubber hand and brushing real and rubber hands synchronously or asynchronously. Perceived finger location was measured before and after stimulation. Hand temperature was taken before and during stimulation. Subjective strength of the illusion was assessed by a questionnaire.
Across groups, the RHI was stronger during synchronous stimulation, indicated by self-report and proprioceptive drift. Patients reported a stronger RHI than controls. Self-reported strength of RHI was associated with schizotypy in controls Proprioceptive drift was larger in patients, but only following synchronous stimulation. Further, we observed stimulation-dependent changes in skin temperature. During right hand stimulation, temperature dropped in the stimulated hand and rose in the unstimulated hand. Interestingly, induction of RHI led to an out-of-body experience in one patient, linking body disownership and psychotic experiences.
The RHI is quantitatively and qualitatively stronger in schizophrenia. These findings suggest that patients have a more flexible body representation and weakened sense of self, and potentially indicate abnormalities in temporo-parietal networks implicated in body ownership. Further, results suggest that these body ownership disturbances might be at the heart of a subset of the pathognomonic delusions of passivity.
PMCID: PMC3205058  PMID: 22073126
4.  Bodily pleasure matters: velocity of touch modulates body ownership during the rubber hand illusion 
The sense of body ownership represents a fundamental aspect of our self-consciousness. Influential experimental paradigms, such as the rubber hand illusion (RHI), in which a seen rubber hand is experienced as part of one's body when one's own unseen hand receives congruent tactile stimulation, have extensively examined the role of exteroceptive, multisensory integration on body ownership. However, remarkably, despite the more general current interest in the nature and role of interoception in emotion and consciousness, no study has investigated how the illusion may be affected by interoceptive bodily signals, such as affective touch. Here, we recruited 52 healthy, adult participants and we investigated for the first time, whether applying slow velocity, light tactile stimuli, known to elicit interoceptive feelings of pleasantness, would influence the illusion more than faster, emotionally-neutral, tactile stimuli. We also examined whether seeing another person's hand vs. a rubber hand would reduce the illusion in slow vs. fast stroking conditions, as interoceptive signals are used to represent one's own body from within and it is unclear how they would be integrated with visual signals from another person's hand. We found that slow velocity touch was perceived as more pleasant and it produced higher levels of subjective embodiment during the RHI compared with fast touch. Moreover, this effect applied irrespective of whether the seen hand was a rubber or a confederate's hand. These findings provide support for the idea that affective touch, and more generally interoception, may have a unique contribution to the sense of body ownership, and by implication to our embodied psychological “self.”
PMCID: PMC3792699  PMID: 24115938
rubber hand illusion; pleasant touch; interoception; body ownership; embodiment
5.  Rubber hand illusion induced by touching the face ipsilaterally to a deprived hand: evidence for plastic “somatotopic” remapping in tetraplegics 
Background: Studies in animals and humans indicate that the interruption of body-brain connections following spinal cord injury (SCI) leads to plastic cerebral reorganization.
Objective: To explore whether inducing the Rubber Hand Illusion (RHI) via synchronous multisensory visuo-tactile bodily stimulation may reveal any perceptual correlates of plastic remapping in SCI.
Methods: In 16 paraplegic, 16 tetraplegic and 16 healthy participants we explored whether RHI may be induced by tactile stimuli involving not only the left hand but also the left hemi-face. Touching the participants actual hand or face was either synchronous or asynchronous with tactile stimuli seen on a rubber hand. We assessed two components of the illusion, namely perceived changes in the real hand in space (indexed by proprioceptive drift) and ownership of the rubber hand (indexed by subjective responses to an ad-hoc questionnaire).
Results: Proprioceptive drift and ownership were found in the healthy group only in the condition where the left real and fake hand were touched simultaneously. In contrast, no drift was found in the SCI patients who, however, showed ownership after both synchronous and asynchronous hand stroking. Importantly, only tetraplegics showed the effect also after synchronous face stroking.
Conclusions: RHI may reveal plastic phenomena in SCI. In hand representation-deprived tetraplegics, stimuli on the face (represented contiguously in the somatic and motor systems), drive the sense of hand ownership. This hand-face remapping phenomenon may be useful for restoring a sense of self in massively deprived individuals.
PMCID: PMC4050649  PMID: 24959128
spinal cord injury; rubber hand illusion; somatosensory plasticity; body representation; tetraplegia; face-hand remapping
6.  Proprioceptive drift in the rubber hand illusion is intensified following 1 Hz TMS of the left EBA 
The rubber hand illusion (RHI) is a paradigm used to induce an illusory feeling of owning a dummy hand through congruent multisensory stimulation. Thus, it can grant insights into how our brain represents our body as our own. Recent research has demonstrated an involvement of the extrastriate body area (EBA), an area of the brain that is typically implicated in the perception of non-face body parts, in illusory body ownership. In this experiment, we sought causal evidence for the involvement of the EBA in the RHI. Sixteen participants took part in a sham controlled, 1 Hz repetitive transcranial magnetic stimulation (rTMS) experiment. Participants received (RHI condition) or asynchronous (control) stroking and were asked to report the perceived location of their real hand, as well as the intensity and the temporal onset of experienced ownership of the dummy hand. Following rTMS of the left EBA, participants misjudged their real hand’s location significantly more toward the dummy hand during the RHI than after sham stimulation. This difference in “proprioceptive drift” provides the first causal evidence that the EBA is involved in the RHI and subsequently in body representation and further supports the view that the EBA is necessary for multimodal integration.
PMCID: PMC4045244  PMID: 24926247
rubber hand illusion; transcranial magnetic stimulation; extrastriate body area; body representation; proprioceptive drift
7.  The Human Touch: Skin Temperature during the Rubber Hand Illusion in Manual and Automated Stroking Procedures 
PLoS ONE  2013;8(11):e80688.
A difference in skin temperature between the hands has been identified as a physiological correlate of the rubber hand illusion (RHI). The RHI is an illusion of body ownership, where participants perceive body ownership over a rubber hand if they see it being stroked in synchrony with their own occluded hand. The current study set out to replicate this result, i.e., psychologically induced cooling of the stimulated hand using an automated stroking paradigm, where stimulation was delivered by a robot arm (PHANToMTM force-feedback device). After we found no evidence for hand cooling in two experiments using this automated procedure, we reverted to a manual stroking paradigm, which is closer to the one employed in the study that first produced this effect. With this procedure, we observed a relative cooling of the stimulated hand in both the experimental and the control condition. The subjective experience of ownership, as rated by the participants, by contrast, was strictly linked to synchronous stroking in all three experiments. This implies that hand-cooling is not a strict correlate of the subjective feeling of hand ownership in the RHI. Factors associated with the differences between the two designs (differences in pressure of tactile stimulation, presence of another person) that were thus far considered irrelevant to the RHI appear to play a role in bringing about this temperature effect.
PMCID: PMC3832597  PMID: 24260454
8.  Neural Correlates of the Rubber Hand Illusion in Amputees: A report of Two Cases 
Neurocase  2013;20(4):407-420.
One of the current challenges in the field of advanced prosthetics is the development of artificial limbs that provide the user with detailed sensory feedback. Sensory feedback from our limbs is not only important for proprioceptive awareness and motor control, but also essential for providing us with a feeling of ownership or simply put, the sensation that our limbs actually belong to ourselves. The strong link between sensory feedback and ownership has been repeatedly demonstrated with the so-called rubber hand illusion (RHI), during which individuals are induced with the illusory sensation that an artificial hand is their own. In healthy participants, this occurs via integration of visual and tactile signals, which is primarily supported by multisensory regions in premotor and intraparietal cortices. Here, we describe a functional magnetic resonance imaging (fMRI) study with two upper limb amputees, showing for the first time that the same brain regions underlie ownership sensations of an artificial hand in this population. Albeit preliminary, these findings are interesting from both a theoretical as well as a clinical point of view. From a theoretical perspective, they imply that even years after the amputation, a few seconds of synchronous visuotactile stimulation are sufficient to activate hand-centered multisensory integration mechanisms. From a clinical perspective, they show that a very basic sensation of touch from an artificial hand can be obtained by simple but precisely targeted stimulation of the stump, and suggest that a similar mechanism implemented in prosthetic hands would greatly facilitate ownership sensations and in turn, acceptance of the prosthesis.
PMCID: PMC3998094  PMID: 23682688
Rubber hand illusion; Amputees; Multisensory integration; fMRI; Prosthetics
9.  Pleasant touch moderates the subjective but not objective aspects of body perception 
Un-myelinated C tactile afferents (CT afferents) are a key finding in affective touch. These fibers, which activate in response to a caress-like touch to hairy skin (CT afferents are not found in palm skin), may have more in common with interoceptive systems encoding body ownership, than afferent systems processing other tactile stimuli. We tested whether subjective embodiment of a rubber hand (measured through questionnaire items) was increased when tactile stimulation was applied to the back of the hand at a rate optimal for CT afferents (3 cm/s) vs. stimulation of glabrous skin (on the palm of the hand) or at a non-optimal rate (30 cm/s), which should not activate these fibers. We also collected ratings of tactile pleasantness and a measure of perceived limb position, proprioceptive drift, which is mediated by different mechanisms of multisensory integration than those responsible for feelings of ownership. The results of a multiple regression analysis revealed that proprioceptive drift was a significant predictor of subjective strength of the illusion when tactile stimuli were applied to the back of the hand, regardless of stroking speed. This relationship was modified by pleasantness, with higher ratings when stimulation was applied to the back of the hand at the slower vs. faster stroking speed. Pleasantness was also a unique predictor of illusion strength when fast stroking was applied to the palm of the hand. However, there were no conditions under which pleasantness was a significant predictor of drift. Since the illusion was demonstrated at a non-optimal stroking speed an integrative role for CT afferents within the illusion cannot be fully supported. Pleasant touch, however, does moderate the subjective aspects of the rubber hand illusion, which under certain tactile conditions may interact with proprioceptive information about the body or have a unique influence on subjective body perception.
PMCID: PMC3870280  PMID: 24391563
CT afferents; insula; pleasant touch; proprioceptive drift; rubber hand illusion
10.  Perception of Body Ownership Is Driven by Bayesian Sensory Inference 
PLoS ONE  2015;10(2):e0117178.
Recent studies have shown that human perception of body ownership is highly malleable. A well-known example is the rubber hand illusion (RHI) wherein ownership over a dummy hand is experienced, and is generally believed to require synchronized stroking of real and dummy hands. Our goal was to elucidate the computational principles governing this phenomenon. We adopted the Bayesian causal inference model of multisensory perception and applied it to visual, proprioceptive, and tactile stimuli. The model reproduced the RHI, predicted that it can occur without tactile stimulation, and that synchronous stroking would enhance it. Various measures of ownership across two experiments confirmed the predictions: a large percentage of individuals experienced the illusion in the absence of any tactile stimulation, and synchronous stroking strengthened the illusion. Altogether, these findings suggest that perception of body ownership is governed by Bayesian causal inference—i.e., the same rule that appears to govern the perception of outside world.
PMCID: PMC4320053  PMID: 25658822
11.  Rubber Hand Illusion Affects Joint Angle Perception 
PLoS ONE  2014;9(3):e92854.
The Rubber Hand Illusion (RHI) is a well-established experimental paradigm. It has been shown that the RHI can affect hand location estimates, arm and hand motion towards goals, the subjective visual appearance of the own hand, and the feeling of body ownership. Several studies also indicate that the peri-hand space is partially remapped around the rubber hand. Nonetheless, the question remains if and to what extent the RHI can affect the perception of other body parts. In this study we ask if the RHI can alter the perception of the elbow joint. Participants had to adjust an angular representation on a screen according to their proprioceptive perception of their own elbow joint angle. The results show that the RHI does indeed alter the elbow joint estimation, increasing the agreement with the position and orientation of the artificial hand. Thus, the results show that the brain does not only adjust the perception of the hand in body-relative space, but it also modifies the perception of other body parts. In conclusion, we propose that the brain continuously strives to maintain a consistent internal body image and that this image can be influenced by the available sensory information sources, which are mediated and mapped onto each other by means of a postural, kinematic body model.
PMCID: PMC3966820  PMID: 24671172
12.  Rubber Hand Illusion under Delayed Visual Feedback 
PLoS ONE  2009;4(7):e6185.
Rubber hand illusion (RHI) is a subject's illusion of the self-ownership of a rubber hand that was touched synchronously with their own hand. Although previous studies have confirmed that this illusion disappears when the rubber hand was touched asynchronously with the subject's hand, the minimum temporal discrepancy of these two events for attenuation of RHI has not been examined.
Methodology/Principal Findings
In this study, various temporal discrepancies between visual and tactile stimulations were introduced by using a visual feedback delay experimental setup, and RHI effects in each temporal discrepancy condition were systematically tested. The results showed that subjects felt significantly greater RHI effects with temporal discrepancies of less than 300 ms compared with longer temporal discrepancies. The RHI effects on reaching performance (proprioceptive drift) showed similar conditional differences.
Our results first demonstrated that a temporal discrepancy of less than 300 ms between visual stimulation of the rubber hand and tactile stimulation to the subject's own hand is preferable to induce strong sensation of RHI. We suggest that the time window of less than 300 ms is critical for multi-sensory integration processes constituting the self-body image.
PMCID: PMC2702687  PMID: 19587780
13.  Losing One's Hand: Visual-Proprioceptive Conflict Affects Touch Perception 
PLoS ONE  2009;4(9):e6920.
While the sense of bodily ownership has now been widely investigated through the rubber hand illusion (RHI), very little is known about the sense of disownership. It has been hypothesized that the RHI also affects the ownership feelings towards the participant's own hand, as if the rubber hand replaced the participant's actual hand. Somatosensory changes observed in the participants' hand while experiencing the RHI have been taken as evidence for disownership of their real hand. Here we propose a theoretical framework to disambiguate whether such somatosensory changes are to be ascribed to the disownership of the real hand or rather to the anomalous visuo-proprioceptive conflict experienced by the participant during the RHI.
Methodology/Principal Findings
In experiment 1, reaction times (RTs) to tactile stimuli delivered to the participants' hand slowed down following the establishment of the RHI. In experiment 2, the misalignment of visual and proprioceptive inputs was obtained via prismatic displacement, a situation in which ownership of the seen hand was doubtless. This condition slowed down the participants' tactile RTs. Thus, similar effects on touch perception emerged following RHI and prismatic displacement. Both manipulations also induced a proprioceptive drift, toward the fake hand in the first experiment and toward the visual position of the participants' hand in the second experiment.
These findings reveal that somatosensory alterations in the experimental hand resulting from the RHI result from cross-modal mismatch between the seen and felt position of the hand. As such, they are not necessarily a signature of disownership.
PMCID: PMC2732904  PMID: 19738900
14.  The Illusion of Owning a Third Arm 
PLoS ONE  2011;6(2):e17208.
Could it be possible that, in the not-so-distant future, we will be able to reshape the human body so as to have extra limbs? A third arm helping us out with the weekly shopping in the local grocery store, or an extra artificial limb assisting a paralysed person? Here we report a perceptual illusion in which a rubber right hand, placed beside the real hand in full view of the participant, is perceived as a supernumerary limb belonging to the participant's own body. This effect was supported by questionnaire data in conjunction with physiological evidence obtained from skin conductance responses when physically threatening either the rubber hand or the real one. In four well-controlled experiments, we demonstrate the minimal required conditions for the elicitation of this “supernumerary hand illusion”. In the fifth, and final experiment, we show that the illusion reported here is qualitatively different from the traditional rubber hand illusion as it is characterised by less disownership of the real hand and a stronger feeling of having two right hands. These results suggest that the artificial hand ‘borrows’ some of the multisensory processes that represent the real hand, leading to duplication of touch and ownership of two right arms. This work represents a major advance because it challenges the traditional view of the gross morphology of the human body as a fundamental constraint on what we can come to experience as our physical self, by showing that the body representation can easily be updated to incorporate an additional limb.
PMCID: PMC3044173  PMID: 21383847
15.  The Rubber Hand Illusion: Feeling of Ownership and Proprioceptive Drift Do Not Go Hand in Hand 
PLoS ONE  2011;6(6):e21659.
In the Rubber Hand Illusion, the feeling of ownership of a rubber hand displaced from a participant's real occluded hand is evoked by synchronously stroking both hands with paintbrushes. A change of perceived finger location towards the rubber hand (proprioceptive drift) has been reported to correlate with this illusion. To measure the time course of proprioceptive drift during the Rubber Hand Illusion, we regularly interrupted stroking (performed by robot arms) to measure perceived finger location. Measurements were made by projecting a probe dot into the field of view (using a semi-transparent mirror) and asking participants if the dot is to the left or to the right of their invisible hand (Experiment 1) or to adjust the position of the dot to that of their invisible hand (Experiment 2). We varied both the measurement frequency (every 10 s, 40 s, 120 s) and the mode of stroking (synchronous, asynchronous, just vision). Surprisingly, with frequent measurements, proprioceptive drift occurs not only in the synchronous stroking condition but also in the two control conditions (asynchronous stroking, just vision). Proprioceptive drift in the synchronous stroking condition is never higher than in the just vision condition. Only continuous exposure to asynchronous stroking prevents proprioceptive drift and thus replicates the differences in drift reported in the literature. By contrast, complementary subjective ratings (questionnaire) show that the feeling of ownership requires synchronous stroking and is not present in the asynchronous stroking condition. Thus, subjective ratings and drift are dissociated. We conclude that different mechanisms of multisensory integration are responsible for proprioceptive drift and the feeling of ownership. Proprioceptive drift relies on visuoproprioceptive integration alone, a process that is inhibited by asynchronous stroking, the most common control condition in Rubber Hand Illusion experiments. This dissociation implies that conclusions about feelings of ownership cannot be drawn from measuring proprioceptive drift alone.
PMCID: PMC3125296  PMID: 21738756
16.  Fake hands in action: embodiment and control of supernumerary limbs 
Demonstrations that the brain can incorporate a fake limb into our bodily representations when stroked in synchrony with our unseen real hand [(the rubber hand illusion (RHI)] are now commonplace. Such demonstrations highlight the dynamic flexibility of the perceptual body image, but evidence for comparable RHI-sensitive changes in the body schema used for action is less common. Recent evidence from the RHI supports a distinction between bodily representations for perception (body image) and for action (body schema) (Kammers et al. in Neuropsychologia 44:2430–2436, 2006). The current study challenges and extends these findings by demonstrating that active synchronous stroking of a brush not only elicits perceptual embodiment of a fake limb (body image) but also affects subsequent reaching error (body schema). Participants were presented with two moving fake left hands. When only one was synchronous during active touch, ownership was claimed for the synchronous hand only and the accuracy of reaching was consistent with control of the synchronous hand. When both fake hands were synchronous, ownership was claimed over both, but only one was controlled. Thus, it would appear that fake limbs can be incorporated into the body schema as well as the body image, but while multiple limbs can be incorporated into the body image, the body schema can accommodate only one.
PMCID: PMC2895889  PMID: 20012536
Rubber hand illusion; Body schema; Body image; Supernumerary limb illusion; Sensorimotor integration
17.  Does Seeing Ice Really Feel Cold? Visual-Thermal Interaction under an Illusory Body-Ownership 
PLoS ONE  2012;7(11):e47293.
Although visual information seems to affect thermal perception (e.g. red color is associated with heat), previous studies have failed to demonstrate the interaction between visual and thermal senses. However, it has been reported that humans feel an illusory thermal sensation in conjunction with an apparently-thermal visual stimulus placed on a prosthetic hand in the rubber hand illusion (RHI) wherein an individual feels that a prosthetic (rubber) hand belongs to him/her. This study tests the possibility that the ownership of the body surface on which a visual stimulus is placed enhances the likelihood of a visual-thermal interaction. We orthogonally manipulated three variables: induced hand-ownership, visually-presented thermal information, and tactically-presented physical thermal information. Results indicated that the sight of an apparently-thermal object on a rubber hand that is illusorily perceived as one's own hand affects thermal judgments about the object physically touching this hand. This effect was not observed without the RHI. The importance of ownership of a body part that is touched by the visual object on the visual-thermal interaction is discussed.
PMCID: PMC3492393  PMID: 23144814
18.  The Sense of the Body in Individuals with Spinal Cord Injury 
PLoS ONE  2012;7(11):e50757.
Increasing evidence suggests that the basic foundations of the self lie in the brain systems that represent the body. Specific sensorimotor stimulation has been shown to alter the bodily self. However, little is known about how disconnection of the brain from the body affects the phenomenological sense of the body and the self. Spinal cord injury (SCI) patients who exhibit massively reduced somatomotor processes below the lesion in the absence of brain damage are suitable for testing the influence of body signals on two important components of the self–the sense of disembodiment and body ownership. We recruited 30 SCI patients and 16 healthy participants, and evaluated the following parameters: (i) depersonalization symptoms, using the Cambridge Depersonalization Scale (CDS), and (ii) measures of body ownership, as quantified by the rubber hand illusion (RHI) paradigm. We found higher CDS scores in SCI patients, which show increased detachment from their body and internal bodily sensations and decreasing global body ownership with higher lesion level. The RHI paradigm reveals no alterations in the illusory ownership of the hand between SCI patients and controls. Yet, there was no typical proprioceptive drift in SCI patients with intact tactile sensation on the hand, which might be related to cortical reorganization in these patients. These results suggest that disconnection of somatomotor inputs to the brain due to spinal cord lesions resulted in a disturbed sense of an embodied self. Furthermore, plasticity-related cortical changes might influence the dynamics of the bodily self.
PMCID: PMC3510173  PMID: 23209824
19.  Assessing a standardised approach to measuring corticospinal integrity after stroke with DTI☆ 
NeuroImage : Clinical  2013;2:521-533.
The structural integrity of the corticospinal tract (CST) after stroke is closely linked to the degree of motor impairment. Simple and reliable methods of assessing white matter integrity within the CST would facilitate the use of this measure in routine clinical practice. Commonly, diffusion tensor imaging is used to measure voxel-wise fractional anisotropy (FA) in a variety of regions of interest (ROIs) representing the CST. Several methods are currently in use with no consensus about which approach is best. ROIs are usually either the whole CST or the posterior limb of the internal capsule (PLIC). These are created manually on brain images or with reference to an individual's CST determined by tractography. Once the ROI has been defined, the FA can be reported as an absolute measure from the ipsilesional side or as a ratio in comparison to the contralesional side. Both corticospinal tracking and manual ROI definition in individual stroke patients are time consuming and subject to bias. Here, we investigated whether using a CST template derived from healthy volunteers was a feasible method for defining the appropriate ROI within which to measure changes in FA. We reconstructed the CST connecting the primary motor cortex to the ipsilateral pons in 23 age-matched control subjects and 21 stroke patients. An average healthy CST template was created from the 23 control subjects. For each patient, FA values were then calculated for both the template CST and for their own CST. We compared patients' FA metrics between the two tracts by considering four measures (FA in the ipsilesional side, FA in the contralesional side, FA ratio of the ipsilesional side to the contralesional side and FA asymmetry between the two sides) and in two tract-based ROIs (whole tract and tract section traversing the PLIC). There were no significant differences in FA metrics for either method, except for contralesional FA. Furthermore, we found that FA metrics relating to CST damage all correlated with motor ability post-stroke equally well. These results suggest that the healthy CST template could be a surrogate structure for defining tract-based ROIs with which to measure stroke patients' FA metrics, avoiding the necessity for CST tracking in individual patients. CST template-based automated quantification of structural integrity would greatly facilitate implementation of practical clinical applications of diffusion tensor imaging.
•Corticospinal tract (CST) integrity can be assessed using fractional anisotropy.•Defining corticospinal tract ROIs in individual stroke patients is open to bias.•FA-based CST integrity was equivalent when using individual or control CST ROIs.•FA-based CST integrity in either ROI correlated with motor impairment equally well.•Assessment of CST integrity with standard CST ROIs is feasible.
PMCID: PMC3777681  PMID: 24179804
Diffusion tensor imaging; Fractional anisotropy; Corticospinal tract; Stroke; Motor ability
20.  White matter structural connectivity is associated with sensorimotor function in stroke survivors☆ 
NeuroImage : Clinical  2013;2:767-781.
Diffusion tensor imaging (DTI) provides functionally relevant information about white matter structure. Local anatomical connectivity information combined with fractional anisotropy (FA) and mean diffusivity (MD) may predict functional outcomes in stroke survivors. Imaging methods for predicting functional outcomes in stroke survivors are not well established. This work uses DTI to objectively assess the effects of a stroke lesion on white matter structure and sensorimotor function.
A voxel-based approach is introduced to assess a stroke lesion's global impact on motor function. Anatomical T1-weighted and diffusion tensor images of the brain were acquired for nineteen subjects (10 post-stroke and 9 age-matched controls). A manually selected volume of interest was used to alleviate the effects of stroke lesions on image registration. Images from all subjects were registered to the images of the control subject that was anatomically closest to Talairach space. Each subject's transformed image was uniformly seeded for DTI tractography. Each seed was inversely transformed into the individual subject space, where DTI tractography was conducted and then the results were transformed back to the reference space. A voxel-wise connectivity matrix was constructed from the fibers, which was then used to calculate the number of directly and indirectly connected neighbors of each voxel. A novel voxel-wise indirect structural connectivity (VISC) index was computed as the average number of direct connections to a voxel's indirect neighbors. Voxel-based analyses (VBA) were performed to compare VISC, FA, and MD for the detection of lesion-induced changes in sensorimotor function. For each voxel, a t-value was computed from the differences between each stroke brain and the 9 controls. A series of linear regressions was performed between Fugl-Meyer (FM) assessment scores of sensorimotor impairment and each DTI metric's log number of voxels that differed from the control group.
Correlation between the logarithm of the number of significant voxels in the ipsilesional hemisphere and total Fugl-Meyer score was moderate for MD (R2 = 0.512), and greater for VISC (R2 = 0.796) and FA (R2 = 0.674). The slopes of FA (p = 0.0036), VISC (p = 0.0005), and MD (p = 0.0199) versus the total FM score were significant. However, these correlations were driven by the upper extremity motor component of the FM score (VISC: R2 = 0.879) with little influence of the lower extremity motor component (FA: R2 = 0.177).
The results suggest that a voxel-wise metric based on DTI tractography can predict upper extremity sensorimotor function of stroke survivors, and that supraspinal intraconnectivity may have a less dominant role in lower extremity function.
•An intrinsic voxel-based structural connectivity metric is proposed.•The metric enhances the impact of stroke lesions on the distant voxels.•Whole-brain extralesional anatomical connectivity predicts functional outcome.•Functional impact of a lesion is determined by residual anatomical connectivity.•Connectivity to the posterior parietal cortex is a key to sensorimotor function.
PMCID: PMC3777792  PMID: 24179827
DTI, diffusion tensor imaging; FA, fractional anisotropy; FOV, field of view; FM, Fugl-Meyer; LDV, log difference volume; LE, lower extremity; MD, mean diffusivity; TE, echo time; TFIRE, Tactful Functional Imaging Research Environment; TR, repetition time; UE, upper extremity; VISC, voxel-wise indirect structural connectivity; Voxel-wise structural connectivity; Tractography; Diffusion tensor imaging; Stroke; Sensorimotor function; Lesion analysis
21.  Exploring the Impact of Ketamine on the Experience of Illusory Body Ownership 
Biological Psychiatry  2011;69(1):35-41.
Our sense of body ownership is profound and familiar, yet it may be misleading. In the rubber-hand illusion, synchronous tactile and visual stimulation lead to the experience that a rubber hand is actually one's own. This illusion is stronger in schizophrenia. Given the evidence that ketamine, a noncompetitive N-methyl-D-aspartate antagonist reproduces symptoms of schizophrenia, we sought to determine whether the rubber-hand illusion is augmented by ketamine.
We studied 15 healthy volunteers in a within-subjects placebo-controlled study. All volunteers carried out two versions of the rubber-hand task, each under both placebo and ketamine infusions. In one task, they saw a rubber hand being stroked in synchrony with tactile stimulation of their real, hidden hand. In the other, stroking of the real and rubber hands was asynchronous. We recorded subjective changes in sense of ownership, as well as participants' ability to localize their hidden hand.
Ketamine was associated with significant increases in subjective measures of the illusion and in hand mislocalization. Although asynchronous visuotactile stimulation attenuates the strength of the illusion during both placebo and ketamine, there remained a significant illusory effect during asynchronous visuotactile stimulation under ketamine compared with placebo. The strength of the illusion during asynchronous visuotactile stimulation correlated with other subjective effects of the drug.
Ketamine mimics the perturbed sense of body ownership seen in schizophrenia, suggesting that it produces a comparable alteration in integration of information across sensory domains and in the subjective and behavioral consequences of such integration.
PMCID: PMC3025328  PMID: 20947068
Body ownership; ketamine; psychosis; rubber hand
22.  The virtual-hand illusion: effects of impact and threat on perceived ownership and affective resonance 
The rubber hand illusion refers to the observation that participants perceive “body ownership” for a rubber hand if it moves, or is stroked in synchrony with the participant's real (covered) hand. Research indicates that events targeting artificial body parts can trigger affective responses (affective resonance) only with perceived body ownership, while neuroscientific findings suggest affective resonance irrespective of ownership (e.g., when observing other individuals under threat). We hypothesized that this may depend on the severity of the event. We first replicated previous findings that the rubber hand illusion can be extended to virtual hands—the virtual-hand illusion. We then tested whether hand ownership and affective resonance (assessed by galvanic skin conductance) are modulated by the experience of an event that either “impacted” (a ball hitting the hand) or “threatened” (a knife cutting the hand) the virtual hand. Ownership was stronger if the virtual hand moved synchronously with the participant's own hand, but this effect was independent from whether the hand was impacted or threatened. Affective resonance was mediated by ownership however: In the face of mere impact, participants showed more resonance in the synchronous condition (i.e., with perceived ownership) than in the asynchronous condition. In the face of threat, in turn, affective resonance was independent of synchronicity—participants were emotionally involved even if a threat was targeting a hand that they did not perceive as their own. Our findings suggest that perceived body ownership and affective responses to body-related impact or threat can be dissociated and are thus unlikely to represent the same underlying process. We argue that affective reactions to impact are produced in a top-down fashion if the impacted effector is assumed to be part of one's own body, whereas threatening events trigger affective responses more directly in a bottom-up fashion—irrespective of body ownership.
PMCID: PMC3764400  PMID: 24046762
vibrotactile stimulation; rubber hand illusion; virtual hand illusion; body ownership; body awareness; threat; affective responses
23.  Referral of Touch and Ownership between the Hands and the Role of the Somatosensory Cortices 
PLoS ONE  2013;8(1):e52768.
Recent studies have shown that the feeling of body ownership can be fooled by simple visuo-tactile manipulations. Perceptual illusions have been reported in which participants sense phantom touch seen on a rubber hand (rubber hand illusion). While previous studies used homologous limbs for those experiments, we here examined an illusion where people feel phantom touch on a left rubber hand when they see it brushed simultaneously with brushes applied to their right hand. Thus, we investigated a referral of touch from the right to the left hand (across the body midline). Since it is known from animal studies that tactile illusions may alter early sensory processing, we expected a modulation of the primary somatosensory cortex (SI) corresponding to this illusion. Neuromagnetic source imaging of the functional topographic organization in SI showed a shift in left SI, associated with the strength of the referral of touch. Hence, we argue that SI seems to be closely associated with this perceptual illusion. The results suggest that the transfer of tactile information across the body midline could be mediated by neurons with bilateral tactile receptive fields (most likely BA2).
PMCID: PMC3534712  PMID: 23300992
24.  When Right Feels Left: Referral of Touch and Ownership between the Hands 
PLoS ONE  2009;4(9):e6933.
Feeling touch on a body part is paradigmatically considered to require stimulation of tactile afferents from the body part in question, at least in healthy non-synaesthetic individuals. In contrast to this view, we report a perceptual illusion where people experience “phantom touches” on a right rubber hand when they see it brushed simultaneously with brushes applied to their left hand. Such illusory duplication and transfer of touch from the left to the right hand was only elicited when a homologous (i.e., left and right) pair of hands was brushed in synchrony for an extended period of time. This stimulation caused the majority of our participants to perceive the right rubber hand as their own and to sense two distinct touches – one located on the right rubber hand and the other on their left (stimulated) hand. This effect was supported by quantitative subjective reports in the form of questionnaires, behavioral data from a task in which participants pointed to the felt location of their right hand, and physiological evidence obtained by skin conductance responses when threatening the model hand. Our findings suggest that visual information augments subthreshold somatosensory responses in the ipsilateral hemisphere, thus producing a tactile experience from the non-stimulated body part. This finding is important because it reveals a new bilateral multisensory mechanism for tactile perception and limb ownership.
PMCID: PMC2734169  PMID: 19742313
25.  Sleep Dissolves Illusion: Sleep Withstands Learning of Visuo-Tactile-Proprioceptive Integration Induced by Repeated Days of Rubber Hand Illusion Training 
PLoS ONE  2014;9(1):e85734.
Multisensory integration is a key factor in establishing bodily self-consciousness and in adapting humans to novel environments. The rubber hand illusion paradigm, in which humans can immediately perceive illusory ownership to an artificial hand, is a traditional technique for investigating multisensory integration and the feeling of illusory ownership. However, the long-term learning properties of the rubber hand illusion have not been previously investigated. Moreover, although sleep contributes to various aspects of cognition, including learning and memory, its influence on illusory learning of the artificial hand has not yet been assessed. We determined the effects of daily repetitive training and sleep on learning visuo-tactile-proprioceptive sensory integration and illusory ownership in healthy adult participants by using the traditional rubber hand illusion paradigm. Subjective ownership of the rubber hand, proprioceptive drift, and galvanic skin response were measured to assess learning indexes. Subjective ownership was maintained and proprioceptive drift increased with daily training. Proprioceptive drift, but not subjective ownership, was significantly attenuated after sleep. A significantly greater reduction in galvanic skin response was observed after wakefulness compared to after sleep. Our results suggest that although repetitive rubber hand illusion training facilitates multisensory integration and physiological habituation of a multisensory incongruent environment, sleep corrects illusional integration and habituation based on experiences in a multisensory incongruent environment. These findings may increase our understanding of adaptive neural processes to novel environments, specifically, bodily self-consciousness and sleep-dependent neuroplasticity.
PMCID: PMC3894989  PMID: 24465671

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