Synchronous Tactile Stimulation Research Articles

No reduction in motor-evoked potential amplitude during the rubber hand illusion.

In the rubber hand illusion (RHI), touches are applied to a fake hand at the same time as touches are applied to a participant's real hand that is hidden in a congruent position. Synchronous (but not asynchronous) tactile stimulation of the two hands may induce the sensation that the fake hand is the participant's own. As such, the illusion is commonly used to examine the sense of body ownership. Some studies indicate that in addition to the subjective experience of limb ownership reported by participants, the RHI can also reduce corticospinal excitability (e.g., as reflected in motor-evoked potential [MEP] amplitude) and alter parietal-motor cortical connectivity in passive participants. These findings have been taken to support a link between motor cortical processing and the subjective experience of body ownership. In this study, we tried to replicate the reduction in MEP amplitude associated with the RHI and uncover the components of the illusion that might explain these changes. As such, we used single-pulse transcranial magnetic stimulation to probe the excitability of the corticospinal motor system as participants experienced the RHI. Despite participants reporting the presence of the illusion and showing shifts in perceived real hand position towards the fake limb supporting its elicitation, we did not observe any associated reduction in MEP amplitude. We conclude that a reduction in MEP amplitude is not a reliable outcome of the RHI and argue that if such effects do occur, they are unlikely to be large or functionally relevant.

Follow Your Nose: Extended Arm Reach After Pinocchio Illusion in Virtual Reality

In this study, we recreate the Pinocchio Illusion—a bodily illusion whereby the perceived length of one’s nose is extended—in Virtual Reality. Participants (n = 38) self-administered tapping on the tip of the nose of a virtual avatar seen from the first-person perspective (using a hand-held controller) while the nose of the avatar slowly grew with each tap. The stimulating virtual arm and the virtual nose were linked such that while the nose grew the arm extended, and then also grew up to 50%. This produced an extension of the perceived reach of the stimulating arm, and an outward drift in the participants’ real arm. A positive correlation between the extent of the outward drift of the participants’ arm and the perceived reachability of distal objects was observed. These results were found both with synchronous tactile stimulation on the participants’ real nose, and without, but not for control conditions in which the visuomotor synchrony or body schema were violated. These findings open new avenues for hand grasp interactions with virtual objects out of arm’s-reach in immersive setups and are discussed in the context of theories of body ownership, body schema, and touch perception.

Deficits in Sense of Body Ownership, Sensory Processing, and Temporal Perception in Schizophrenia Patients With/Without Auditory Verbal Hallucinations.

It has been claimed that individuals with schizophrenia have difficulty in self-recognition and, consequently, are unable to identify the sources of their sensory perceptions or thoughts, resulting in delusions, hallucinations, and unusual experiences of body ownership. The deficits also contribute to the enhanced rubber hand illusion (RHI; a body perception illusion, induced by synchronous visual and tactile stimulation). Evidence based on RHI paradigms is emerging that auditory information can make an impact on the sense of body ownership, which relies on the process of multisensory inputs and integration. Hence, we assumed that auditory verbal hallucinations (AVHs), as an abnormal auditory perception, could be linked with body ownership, and the RHI paradigm could be conducted in patients with AVHs to explore the underlying mechanisms. In this study, we investigated the performance of patients with/without AVHs in the RHI. We administered the RHI paradigm to 80 patients with schizophrenia (47 with AVHs and 33 without AVHs) and 36 healthy controls. We conducted the experiment under two conditions (synchronous and asynchronous) and evaluated the RHI effects by both objective and subjective measures. Both patient groups experienced the RHI more quickly and strongly than HCs. The RHI effects of patients with AVHs were significantly smaller than those of patients without AVHs. Another important finding was that patients with AVHs did not show a reduction in RHI under asynchronous conditions. These results emphasize the disturbances of the sense of body ownership in schizophrenia patients with/without AVHs and the associations with AVHs. Furthermore, it is suggested that patients with AVHs may have multisensory processing dysfunctions and internal timing deficits.

Hypothesis awareness confounds asynchronous control conditions in indirect measures of the rubber hand illusion.

Reports of changes in experiences of body location and ownership following synchronous tactile and visual stimulation of fake and real hands (rubber hand (RH) effects) are widely attributed to multisensory integration mechanisms. However, existing control methods for subjective report measures (asynchronous stroking and control statements) are confounded by participant hypothesis awareness; the report may reflect response to demand characteristics. Subjective report is often accompanied by indirect (also called ‘objective’ or ‘implicit’) measures. Here, we report tests of expectancies for synchronous ‘illusion’ and asynchronous ‘control’ conditions across two pre-registered studies (n = 140 and n = 45) for two indirect measures: proprioceptive drift (a change in perceived hand location) and skin conductance response (a measure of physiological arousal). Expectancies for synchronous condition measures were greater than for asynchronous conditions in both studies. Differences between synchronous and asynchronous control condition measures are therefore confounded by hypothesis awareness. This means indirect measures of RH effects may reflect compliance, bias and phenomenological control in response to demand characteristics, just as for subjective measures. Valid control measures are required to support claims of a role of multisensory integration for both direct and indirect measures of RH effects.

Investigating the Hand Ownership Illusion With Two Views Merged in.

Researchers investigating virtual/augmented reality have shown humans' marked adaptability, especially regarding our sense of body ownership; their cumulative findings have expanded the concept of what it means to have a body. Herein, we report the hand ownership illusion during “two views merged in.” In our experiment, participants were presented two first-person perspective views of their arm overlapped, one was the live feed from a camera and the other was a playback video of the same situation, slightly shifted toward one side. The relative visibility of these two views and synchrony of tactile stimulation were manipulated. Participants' level of embodiment was evaluated using a questionnaire and proprioceptive drift. The results show that the likelihood of embodying the virtual hand is affected by the relative visibility of the two views and synchrony of the tactile events. We observed especially strong hand ownership of the virtual hand in the context of high virtual hand visibility with synchronous tactile stimulation.

Multisensory Facial Stimulation Implicitly Improves Evaluations of the Goodness of Attractive Others.

It has been well demonstrated that shared multisensory experiences between the self and others can influence the social perception of out-group members. Previous research has shown that the illusion of ownership over a dark-skinned rubber hand or full virtual body generated less negative implicit bias against people with dark skin. However, less is known about how perceived attractiveness difference between self and other affects social perception toward those others after shared multisensory experience. The present study assessed whether shared multisensory experience between the self and attractive others would affect the implicit evaluation of goodness of others. Seventy-three women participated in the study. After the visuotactile multisensory stimulation procedure, participants were administered the Single Category Implicit Association Test (SC-IAT), which presents two attributes (good and bad) and one concept (other). Results showed that the more attractive the faces are, the more positive their implicit evaluation becomes after the synchronous tactile stimulation. This result suggests that shared multisensory experience makes people feel more positive toward others who have positive attribute. This finding suggests that self-other blurring in social contexts might be a compelling factor in evaluating other people positively.

Dynamic Displacement Vector Interacts with Tactile Localization

SummaryLocating a tactile stimulus on the body seems effortless and straightforward. However, the perceived location of a tactile stimulation can differ from its physical location [1, 2, 3]. Tactile mislocalizations can depend on the timing of successive stimulations [2, 4, 5], tactile motion mechanisms [6], or processes that “remap” stimuli from skin locations to external space coordinates [7, 8, 9, 10, 11]. We report six experiments demonstrating that the perception of tactile localization on a static body part is strongly affected by the displacement between the locations of two successive task-irrelevant actions. Participants moved their index finger between two keys. Each keypress triggered synchronous tactile stimulation at a randomized location on the immobilized wrist or forehead. Participants reported the location of the second tactile stimulation relative to the first. The direction of either active finger movements or passive finger displacements biased participants’ tactile orientation judgements (experiment 1). The effect generalized to tactile stimuli delivered to other body sites (experiment 2). Two successive keypresses, by different fingers at distinct locations, reproduced the effect (experiment 3). The effect remained even when the hand that moved was placed far from the tactile stimulation site (experiments 4 and 5). Temporal synchrony within 600 ms between the movement and tactile stimulations was necessary for the effect (experiment 6). Our results indicate that a dynamic displacement vector, defined as the location of one sensorimotor event relative to the one before, plays a strong role in structuring tactile spatial perception.

Dynamic causal modeling of touch-evoked potentials in the rubber hand illusion

The neural substrate of bodily ownership can be disclosed by the rubber hand illusion (RHI); namely, the illusory self-attribution of an artificial hand that is induced by synchronous tactile stimulation of the subject's hand that is hidden from view. Previous studies have pointed to the premotor cortex (PMC) as a pivotal area in such illusions. To investigate the effective connectivity between – and within – sensory and premotor areas involved in bodily perceptions, we used dynamic causal modeling of touch-evoked responses in 13 healthy subjects. Each subject's right hand was stroked while viewing their own hand (“REAL”), or an artificial hand presented in an anatomically plausible (“CONGRUENT”) or implausible (“INCONGRUENT”) position. Bayesian model comparison revealed strong evidence for a differential involvement of the PMC in the generation of touch-evoked responses under the three conditions, confirming a crucial role of PMC in bodily self-attribution. In brief, the extrinsic (forward) connection from left occipital cortex to left PMC was stronger for CONGRUENT and INCONGRUENT as compared to REAL, reflecting the augmentation of bottom-up visual input when multisensory integration is challenged. Crucially, intrinsic connectivity in the primary somatosensory cortex (S1) was attenuated in the CONGRUENT condition, during the illusory percept. These findings support predictive coding models of the functional architecture of multisensory integration (and attenuation) in bodily perceptual experience.

Sensory processing and the rubber hand illusion--an evoked potentials study.

The rubber hand illusion (RHI) paradigm--in which illusory bodily ownership is induced by synchronous tactile stimulation of a participant's (hidden) hand and a (visible) surrogate--allows one to investigate how the brain resolves conflicting multisensory evidence during perceptual inference. To identify the functional anatomy of the RHI, we used multichannel EEG, acquired under three conditions of tactile stimulation. Evoked potentials were averaged from EEG signals registered to the timing of brushstrokes to the participant's hand. The participant's hand was stroked either in the absence of an artificial hand (REAL) or synchronously with an artificial hand, which either lay in an anatomically plausible (CONGRUENT) or impossible (INCONGRUENT) position. The illusion was reliably elicited in the CONGRUENT condition. For right-hand stimulation, significant differences between conditions emerged at the sensor level around 55 msec after the brushstroke at left frontal and right parietal electrodes. Response amplitudes were smaller for illusory (CONGRUENT) compared with nonillusory (INCONGRUENT and REAL) conditions in the contralateral perirolandic region (pre- and postcentral gyri), superior and inferior parietal lobule, whereas veridical perception of the artificial hand (INCONGRUENT) amplified responses at a scalp region overlying the contralateral postcentral gyrus and inferior parietal lobule compared with the remaining two conditions. Left-hand stimulation produced similar contralateral patterns. These results are consistent with predictive coding models of multisensory integration and may reflect the attenuation of somatosensory precision that is required to resolve perceptual hypotheses about conflicting multisensory input.

Crossing the Hands Increases Illusory Self-Touch

Manipulation of hand posture, such as crossing the hands, has been frequently used to study how the body and its immediately surrounding space are represented in the brain. Abundant data show that crossed arms posture impairs remapping of tactile stimuli from somatotopic to external space reference frame and deteriorates performance on several tactile processing tasks. Here we investigated how impaired tactile remapping affects the illusory self-touch, induced by the non-visual variant of the rubber hand illusion (RHI) paradigm. In this paradigm blindfolded participants (Experiment 1) had their hands either uncrossed or crossed over the body midline. The strength of illusory self-touch was measured with questionnaire ratings and proprioceptive drift. Our results showed that, during synchronous tactile stimulation, the strength of illusory self-touch increased when hands were crossed compared to the uncrossed posture. Follow-up experiments showed that the increase in illusion strength was not related to unfamiliar hand position (Experiment 2) and that it was equally strengthened regardless of where in the peripersonal space the hands were crossed (Experiment 3). However, while the boosting effect of crossing the hands was evident from subjective ratings, the proprioceptive drift was not modulated by crossed posture. Finally, in contrast to the illusion increase in the non-visual RHI, the crossed hand postures did not alter illusory ownership or proprioceptive drift in the classical, visuo-tactile version of RHI (Experiment 4). We argue that the increase in illusory self-touch is related to misalignment of somatotopic and external reference frames and consequently inadequate tactile-proprioceptive integration, leading to re-weighting of the tactile and proprioceptive signals.The present study not only shows that illusory self-touch can be induced by crossing the hands, but importantly, that this posture is associated with a stronger illusion.

Full body illusion is associated with widespread skin temperature reduction

A central feature of our consciousness is the experience of the self as a unified entity residing in a physical body, termed bodily self-consciousness. This phenomenon includes aspects such as the sense of owning a body (also known as body ownership) and has been suggested to arise from the integration of sensory signals from the body. Several studies have shown that temporally synchronous tactile stimulation of the real body and visual stimulation of a fake or virtual body can induce changes in bodily self-consciousness, typically resulting in a sense of illusory ownership over the fake body. The present study assessed the effect of anatomical congruency of visuo-tactile stimulation on bodily self-consciousness. A virtual body was presented and temporally synchronous visuo-tactile stroking was applied simultaneously to the participants' body and to the virtual body. We manipulated the anatomical locations of the visuo-tactile stroking (i.e., on the back, on the leg), resulting in congruent stroking (stroking was felt and seen on the back or the leg) or incongruent stroking (i.e., stroking was felt on the leg and seen on the back). We measured self-identification with the virtual body and self-location as well as skin temperature. Illusory self-identification with the avatar as well as changes in self-location were experienced in the congruent stroking conditions. Participants showed a decrease in skin temperature across several body locations during congruent stimulation. These data establish that the full-body illusion (FBI) alters bodily self-consciousness and instigates widespread physiological changes in the participant's body.

Drift and ownership toward a distant virtual body

In body ownership illusions participants feel that a mannequin or virtual body (VB) is their own. Earlier results suggest that body ownership over a body seen from behind in extra personal space is possible when the surrogate body is visually stroked and tapped on its back, while spatially and temporal synchronous tactile stimulation is applied to the participant's back. This result has been disputed with the claim that the results can be explained by self-recognition rather than somatic body ownership. We carried out an experiment with 30 participants in a between-groups design. They all saw the back of a VB 1.2 m in front, that moved in real-time determined by upper body motion capture. All felt tactile stimulation on their back, and for 15 of them this was spatially and temporally synchronous with stimulation that they saw on the back of the VB, but asynchronous for the other 15. After 3 min a revolving fan above the VB descended and stopped at the position of the VB neck. A questionnaire assessed referral of touch to the VB, body ownership, the illusion of drifting forwards toward the VB, and the VB drifting backwards. Heart rate deceleration (HRD) and the amount of head movement during the threat period were used to assess the response to the threat from the fan. Results showed that although referral of touch was significantly greater in the synchronous condition than the asynchronous, there were no other differences between the conditions. However, a further multivariate analysis revealed that in the visuotactile synchronous condition HRD and head movement increased with the illusion of forward drift and decreased with backwards drift. Body ownership contributed positively to these drift sensations. Our conclusion is that the setup results in a contradiction—somatic feelings associated with a distant body—that the brain attempts to resolve by generating drift illusions that would make the two bodies coincide.

Exploring the Impact of Ketamine on the Experience of Illusory Body Ownership

BackgroundOur 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.MethodsWe 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.ResultsKetamine 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.ConclusionsKetamine 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.

Inducing a virtual hand ownership illusion through a brain–computer interface

The apparently stable brain representation of our bodies is easily challenged. We have recently shown that the illusion of ownership of a three-dimensional virtual hand can be evoked through synchronous tactile stimulation of a person's hidden real hand and that of the virtual hand. This reproduces the well-known rubber-hand illusion, but in virtual reality. Here we show that some aspects of the illusion can also occur through motor imagery used to control movements of a virtual hand. When movements of the virtual hand followed motor imagery, the illusion of ownership of the virtual hand was evoked and muscle activity measured through electromyogram correlated with movements of the virtual arm. Using virtual bodies has a great potential in the fields of physical and neural rehabilitation, making the understanding of ownership of a virtual body highly relevant.

Referral of sensation to an advanced humanoid robotic hand prosthesis

Hand prostheses that are currently available on the market are used by amputees to only a limited extent, partly because of lack of sensory feedback from the artificial hand. We report a pilot study that showed how amputees can experience a robot-like advanced hand prosthesis as part of their own body. We induced a perceptual illusion by which touch applied to the stump of the arm was experienced from the artificial hand. This illusion was elicited by applying synchronous tactile stimulation to the hidden amputation stump and the robotic hand prosthesis in full view. In five people who had had upper limb amputations this stimulation caused referral touch sensation from the stump to the artificial hand, and the prosthesis was experienced more like a real hand. We also showed that this illusion can work when the amputee controls the movements of the artificial hand by recordings of the arm muscle activity with electromyograms. These observations indicate that the previously described "rubber hand illusion" is also valid for an advanced hand prosthesis, even when it has a robotic-like appearance.

Improvement of sensorimotor functions in old age by passive sensory stimulation

Sensorimotor functions decrease in old age. The well-documented loss of tactile acuity in elderly is accompanied by deterioration ofhaptic performance and fine manipulative movements. Physical training and exercise can maintain sensorimotor fitness into high age. However, regular schedules of training require discipline and physical fitness. We here present an alternative interventional paradigm to enhance tactile, haptic, and fine motor performance based on passive, sensory stimulation by means of tactile coactivation. This approach is based on patterned, synchronous tactile stimulation applied to the fingertips for 3 hours. The stimulation drives plastic reorganizational changes in somatosensory cortex that affect perception and behavior: We demonstrate that following 3 hours of coactivation tactile acuity as well as haptic object exploration and fine motor performance are improved for at least 96 hours. Because this kind of intervention does not require active participation or attention of the subjects, we anticipate that coactivation is a prime candidate for future therapeutic interventions in patients with impaired sensorimotor abilities. It can be assumed that the maintenance and restoration of sensorimotor functions can ensure and preserve independence of daily living. Further optimizing of the stimulation protocol can be assumed to strengthen both the range and durability of its efficacy.

Having a body versus moving your body: How agency structures body-ownership

We investigated how motor agency in the voluntary control of body movement influences body awareness. In the Rubber Hand Illusion (RHI), synchronous tactile stimulation of a rubber hand and the participant’s hand leads to a feeling of the rubber hand being incorporated in the participant’s own body. One quantifiable behavioural correlate of the illusion is an induced shift in the perceived location of the participant’s hand towards the rubber hand. Previous studies showed that the induced changes in body awareness are local and fragmented: the proprioceptive drift is largely restricted to the stimulated finger. In the present study, we investigated whether active and passive movements, rather than tactile stimulation, would lead to similarly fragmented body awareness. Participants watched a projected image of their hand under three conditions: active finger movement, passive finger movement, and tactile stimulation. Visual feedback was either synchronous or asynchronous with respect to stimulation of the hand. A significant overall RHI, defined as greater drifts following synchronous than asynchronous stimulation, was found in all cases. However, the distribution of the RHI across stimulated and non-stimulated fingers depended on the kind of stimulation. Localised proprioceptive drifts, specific to the stimulated finger, were found for tactile and passive stimulation. Conversely, during active movement of a single digit, the proprioceptive drifts were not localised to that digit, but were spread across the whole hand. Whereas a purely proprioceptive sense of body-ownership is local and fragmented, the motor sense of agency integrates distinct body-parts into a coherent, unified awareness of the body.

Tactile "capture" of audition.

Previous research has demonstrated that the localization of auditory or tactile stimuli can be biased by the simultaneous presentation of a visual stimulus from a different spatial position. We investigated whether auditory localization judgments could also be affected by the presentation of spatially displaced tactile stimuli, using a procedure designed to reveal perceptual interactions across modalities. Participants made left-right discrimination responses regarding the perceived location of sounds, which were presented either in isolation or together with tactile stimulation to the fingertips. The results demonstrate that the apparent location of a sound can be biased toward tactile stimulation when it is synchronous, but not when it is asynchronous, with the auditory event. Directing attention to the tactile modality did not increase the bias of sound localization toward synchronous tactile stimulation. These results provide the first demonstration of the tactile capture of audition.

Activity patterns of human somatosensory cortex adapt dynamically to stimulus properties.

Long-term synchronous tactile stimulation of two sites of the body results in integrated, overlapping cortical representations whereas asynchronous stimulation leads to segregated representations. To investigate the cortical capacity to adapt dynamically to stimulation properties 22 subjects were stimulated at digits 1, 3 and 5 of both hands in either random or fixed order. Changes in the functional organization of the somatosensory cortex were inferred by neuromagnetic source analysis based on somatosensory evoked magnetic fields. Compared to the stimulation in random sequence, the stimulation in fixed order revealed a reduction in distance between the cortical representation of D1 and D3. We conclude that the pattern of activation in the somatosensory cortex adapts dynamically to the spatio-temporal characteristics of the stimuli.

Differential Activation in Somatosensory Cortex for Different Discrimination Tasks

Maps of the body surface in somatosensory cortex have been shown to be highly plastic, altering their configuration in response to changes in use of body parts. The current study investigated alterations in the functional organization of the human somatosensory cortex resulting from massed practice. Over a period of 4 weeks, subjects were given synchronous tactile stimulation of thumb (D1) and little finger (D5) for 1 hr/d. They had to identify the orientation of the stimuli. Neuroelectric source localization based on high-resolution EEG revealed that, when subjects received passive tactile stimulation of D1 or D5, the representations of the fingers in primary somatosensory cortex were closer together after training than before. There was also an apparently correlative tendency to anomalously mislocalize near-threshold tactile stimuli equally to the distant finger costimulated during training rather than preferentially to the finger nearest to the finger stimulated in a post-training test. However, when the stimulus discrimination had to be made, neuroelectric source imaging revealed that the digital representations of D1 and D5 were further apart after training than before. Thus, the same series of prolonged repetitive stimulations produced two different opposite effects on the spatial relationship of the cortical representations of the digits, suggesting that differential activation in the same region of somatosensory cortex is specific to different tasks.