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The classical body ownership illusion, such as the rubber hand illusion, is achieved through appropriate proprioceptive displacement within temporal and spatial constraints that do not exceed the limits of proprioceptive flexibility. In the 2023 Best Illusion of the Year Contest, we introduced Buddha's ear illusion (BEI), which creates the illusion of owning a dramatically deformed earlobe through immediate visuotactile stimulation and seemingly challenges classical proprioceptive boundaries. The laboratory experiment examined the mechanics of this illusion, revealing a significant interaction between tactile earlobe pulling and visual miming that contributed to the enhanced perception of earlobe stretch. Importantly, 88% of the participants confirmed the illusory earlobe stretch (a rating of +4 or higher on a 7-point scale). More than half reported an earlobe descent of >10 cm within a 10-s visuotactile stimulation. The findings suggest that BEI operates on a distinct principle separate from proprioceptive modulation in contrast to classical ownership illusions.

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Previous studies have examined the rubber hand illusion with finger lengthening, but there is limited research on finger widening. This suggests a strong cognitive bias toward the illusory expansion of the finger in a distal direction rather than lateral. To test this, we compared the illusory deformability of the finger in the distal and lateral directions through the generation of illusory finger deformation using a double-touch operation, referred to as the numbness illusion. Our results showed that perceived distal distortion was wholly superior to perceived lateral distortion in terms of sense of ownership ratings. Moreover, the extent of the perceived deformation was greater in the distal than lateral direction, supporting our hypothesis that there is a distal bias. We suggest that this preference may be because the presence of multiple joints is required to create illusory deformation in the target direction.

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Embodiment is a complex concept related to the subjective perception of an object as it belongs to its own body. In general, this construct has been evaluated by means of questionnaires, but validation studies in other cultures and limitations related with barriers of language received little attention. The purpose of the present investigation was twofold: to validate the factorial structure of embodiment questionnaire (EQ) and to construct a pictographic scale (PAE) to measure embodiment without relapse verbal representations. In the first experiment, 136 participants underwent a Rubber Hand Illusion (RHI) procedure following both congruent and incongruent (control) visuo-tactile stimulations. Then, they evaluated embodiment illusion in EQ using a Likert-type scale to rate their agreement or disagreement with 27 statements and with a pictographic scale designed to assess their subjective experience of the illusion. Principal components analysis in EQ scores identified four components that emerged in both conditions: Embodiment, Disembodiment, Affect and Deafference. PAE scale was highly correlated with embodiment factor and can differentiate between conditions. In a second experiment, 30 participants underwent the RHI procedure, and they were assessed using PAE and proprioceptive drift. Results indicate a high positive correlation between PAE and post-illusion drift score. These results provide evidence about the consistency of the factorial structure of EQ across cultures, and we also provide a new pictographic tool that allows quick measurement of embodiment overcoming language barriers.

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In the rubber hand illusion (RHI), participants see a fake hand touched synchronously with their real hand, which is hidden from view. The three-way interaction between vision, touch, and proprioception induces the sensation that the dummy hand belongs to oneself (i.e., subjective embodiment) and the illusory displacement of the real hand towards the fake one (i.e., proprioceptive drift). In the literature, there are mixed results (some positive and some null) regarding the existence of a relationship between subjective embodiment and proprioceptive drift. We conducted a Bayesian meta-analysis to tackle this issue quantitatively. Evidence strongly favours the presence of a correlation between subjective embodiment and proprioceptive drift, supporting the model proposed by Botvinick and Cohen in 1998. However, the correlation is around .35, a value suggesting that the two indices capture different facets of the RHI. This result clarifies the association between the illusory effects produced by the RHI and may be helpful for designing studies having appropriate statistical power.

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A growing body of research has shown that a unilateral alteration in the sense of limb ownership is associated with the cooling of a limb's temperature. However, the recent emergence of contradictory results calls into question the existence of a relationship between this physiological reaction and the sense of body ownership. In the light of evidence that the malleability of the sense of hand ownership differs based on the preferential motor use of the hand to which the illusion is applied, one might observe the same lateralised pattern in the skin temperature cooling. In particular, if skin temperature change is a signature of body ownership, we expected a stronger illusion and reduction in skin temperature when altering ownership alteration of the left hand compared to the right hand in dextral individuals. To test this hypothesis, we selectively perturbated body ownership of the left or right hand in 24 healthy participants in different experimental sessions using the Mirror-Box Illusion (MBI) paradigm. Participants were asked to tap synchronously or asynchronously at a constant rhythm with their left and right index fingers against two parallel mirrors while looking at their reflected right/left hand. Skin temperature was measured before and after each MBI application, and explicit judgments of ownership and proprioceptive drift were collected. The results showed a consistent cooling of the hand's temperature only when the illusion was performed on the left hand. Proprioceptive drift exhibited the same pattern. In contrast, the explicit judgment of ownership of the reflected hand was similar across the two hands. These data provide evidence in favor of a specific laterality effect of the physiological response to an induced alteration of body part ownership. Moreover, they highlight the possibility of a direct link between proprioception and skin temperature.

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We explored hand ownership in teenagers with unilateral cerebral palsy (UCP) compared with typically developing teenagers. Eighteen participants with UCP and 16 control teenagers participated. We used the rubber hand illusion to test hand ownership (HO). Both affected/non-affected hands (UCP) and dominant/non-dominant hands (controls) were tested during synchronous and asynchronous strokes. HO was assessed by measuring the proprioceptive drift toward the fake hand (as a percentage of arm length) and conducting a questionnaire on subjective HO. Both groups had significantly higher proprioceptive drift in the synchronous stroking condition for both hands. Teenagers with UCP showed a significantly higher proprioceptive drift when comparing their paretic hand (median 3.4% arm length) with the non-dominant hand of the controls (median 1.7% arm length). The questionnaires showed that synchronous versus asynchronous stroking generated a robust change in subjective HO in the control teenagers, but not in the teenagers with UCP. Teenagers with UCP have an altered sense of HO and a distorted subjective experience of HO that may arise from the early dysfunction of complex sensory-motor integration related to their brain lesions. HO may influence motor impairment and prove to be a target for early intervention.

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Badminton players have a plastic modification of their arm representation in the brain due to the prolonged use of their racket. However, it is not known whether their arm representation can be altered through short-term visuotactile integration. The neural representation of the body is easily altered when multiple sensory signals are integrated in the brain. One of the most popular experimental paradigms for investigating this phenomenon is the "rubber hand illusion." This study was designed to investigate the effect of prolonged use of a racket on the modulation of arm representation during the rubber hand illusion in badminton players. When badminton players hold the racket, their badminton experience in years is negatively correlated with the magnitude of the rubber hand illusion. This finding suggests that tool embodiment obtained by the prolonged use of the badminton racket is less likely to be disturbed when holding the racket.

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Although the illusion that the mirror image of a hand or limb could be recognized as a part of one's body behind the mirror, the effect of adding tactile stimulation to this illusion remains unknown. We, therefore, examined how the timing of tactile stimulation affects the induction of body ownership on the mirror image. Twenty-one healthy, right-handed participants (mean age = 23.0 ± 1.0 years, no medical history of neurological and/or psychiatric disorders) were enrolled and a crossover design was adopted in this study. Participants' right and left hands were placed on the front and back sides of the mirror, respectively, then they were asked to keep looking at their right hand in the mirror. All participants experienced two experiments; one was with tactile stimulation that was synchronized with the movement of a mirror image (synchronous condition), and the other one was with tactile stimulation that was not synchronized (asynchronous condition). The qualitative degree of body ownership for the mirrored hand was evaluated by a questionnaire. Proprioceptive drift (PD), an illusory shift of the felt position of the real hand toward the mirrored hand was used for quantitative evaluation of body ownership and measured at "baseline," "immediately after stimulation," "2 min after stimulation," and "4 min after stimulation." The results of the questionnaire revealed that some items of body ownership rating were higher in the synchronous condition than in the asynchronous condition (p < 0.05). We found that PD occurred from immediately after to 4 min after stimulation in both conditions (p < 0.01) and there was no difference in the results between the conditions. From the dissociation of these results, we interpreted that body ownership could be elicited by different mechanisms depending on the task demand. Our results may contribute to the understanding of the multisensory integration mechanism of visual and tactile stimulation during mirror illusion induction.

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It is often claimed that the human self consists of perceived body ownership and agency, which are commonly assessed through explicit ownership and agency judgments and implicit measures, like proprioceptive drift, skin conductance responses, and intentional binding effects. Bottom-up multisensory integration and top-down modulation were predicted to be important for ownership and agency. In previous studies, cognitive load was revealed to affect the sense of agency in a top-down fashion, but its effect on ownership has not been fully investigated, not even its possibly different effect on explicit and implicit measures. Here we used cognitive load (small vs. large sets in a working-memory task) to disentangle explicit and implicit measures of ownership and agency in a task inducing the virtual hand illusion (VHI; stronger perceived ownership and agency over a virtual hand if it moves in synchrony with one's real hand). Results showed similar patterns for ownership and agency - both ownership and agency were affected by cognitive load, and importantly in the explicit measures, higher load increased the effect of synchrony (i.e., the VHI), but in implicit measures, higher load reduced indications of both ownership and agency. Hence, the load manipulation was selective with regard to the explicit versus implicit nature of the measure but not with respect to the measure's content. This provides strong evidence that explicit and implicit measures of both ownership and agency rely on at least partly separable informational sources, while the difference between ownership and agency as such does not seem to play a major role.

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The sense of self is a complex phenomenon, comprising various sensations of bodily self-consciousness. Interestingly, the experience of possessing a body - 'embodiment' - and locating the body within space may be modulated by the Rubber Hand Illusion (RHI). Measures of the RHI include proprioceptive drift (PD), the extent to which the hand is mis-localised towards the rubber hand, and subjective questionnaires. Although these measures often correlate, research from the visual RHI suggests that they reflect separate underlying processes. We investigated whether increasing the duration of tactile stimulation would affect PD and questionnaires differently during the somatic RHI. Participants experienced 30 s, 2 min, or 5 min of synchronous or asynchronous tactile stimulation. Increasing duration affected only PD, with increased drift following 5 min vs 30 s of stimulation. Our findings suggest that PD and questionnaires are not proxies for one another, but reflect separate underlying processes of the somatic RHI.

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BACKGROUND: The feeling of body ownership relies on the binding of multisensory body-related signals. Various sensory abnormalities have been described in Parkinson's disease (PD). OBJECTIVE: To assess the rubber hand illusion (RHI) in patients with PD (PwPD) and age-matched healthy controls (CTRL). To evaluate the influence of the dopaminergic system in a PwPD subgroup OFF medication. METHODS: The RHI paradigm was applied to 42 PwPD and 48 CTRL. In this experimental setup, stroking a visible plastic hand simultaneously with the covered real hand elicits the feeling of ownership over the seen hand. Asynchronous stroking served as a control condition. Proprioceptive bias and an illusion score based on a questionnaire were used as measures of the RHI. Seventeen PwPD additionally underwent the experiments "OFF medication". RESULTS: Compared to CTRL, PwPD showed higher proprioceptive bias independent of the stroking condition (p = 0.015), and had higher illusion scores in the asynchronous condition (p < 0.05). In PwPD, there were no significant differences between ON- and OFF-medication state. CONCLUSION: In PwPD, responses to the RHI are less specific with respect to the degree of synchronicity of brushstrokes. This might be attributed to a less stable body representation, internal "noise" during multisensory integration, or a blur of temporal discrimination in PD. The fact that RHI measures did not differ between ON- and OFF-medication states indicates an involvement of non-dopaminergic transmitter systems in this finding.

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Virtual reality (VR) is effectively used to evoke the mirror illusion, and transcranial direct current stimulation (tDCS) synergistically facilitates this illusion. This study investigated whether a mirror virtual hand illusion (MVHI) induced by an immersive, first-person-perspective, virtual mirror system could be modulated by tDCS of the primary motor cortex. Fourteen healthy adults (average age 21.86 years ±0.47, seven men and seven women) participated in this study, and they experienced VR with and without tDCS-the tDCS and sham conditions, each of which takes ∼30 minutes-on separate days to allow the washout of the tDCS effect. While experiencing VR, the movements of the virtual left hand reflected the flexion and extension of the real right hand. Subsequently, electroencephalogram was recorded, the magnitude of the proprioceptive shift was measured, and the participants provided responses to a questionnaire regarding hand ownership. A significant difference in the proprioceptive shift was observed between the tDCS and sham conditions. In addition, there was significant suppression of the mu power in Pz, and augmentation of the beta power in the Pz, P4, O1, and O2 channels. The difference in proprioceptive deviation between the two conditions showed significant negative correlation with mu suppression over the left frontal lobe in the tDCS condition. Finally, the question "I felt that the virtual hand was my own hand" received a significantly higher score under the tDCS condition. In short, applying tDCS over the motor cortex facilitates the MVHI by activating the attentional network over the parietal and frontal lobes such that the MVHI induces more proprioceptive drift, which suggests that the combination of VR and tDCS can enhance the immersive effect in VR. This result provides better support for the use of the MVHI paradigm in combination with tDCS for recovery from illnesses such as stroke.

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Spatial action-effect binding denotes the mutual attraction between the perceived position of an effector (e.g., one's own hand) and a distal object that is controlled by this effector. Such spatial binding can be construed as an implicit measure of object ownership, thus the belonging of a controlled object to the own body. The current study investigated how different transformations of hand movements (body-internal action component) into movements of a visual object (body-external action component) affect spatial action-effect binding, and thus implicit object ownership. In brief, participants had to bring a cursor on the computer screen into a predefined target position by moving their occluded hand on a tablet and had to estimate their final hand position. In Experiment 1, we found a significantly lower drift of the proprioceptive position of the hand towards the visual object when hand movements were transformed into laterally inverted cursor movements, rather than cursor movements in the same direction. Experiment 2 showed that this reduction reflected an elimination of spatial action-effect binding in the inverted condition. The results are discussed with respect to the prerequisites for an experience of ownership over artificial, noncorporeal objects. Our results show that predictability of an object movement alone is not a sufficient condition for ownership because, depending on the type of transformation, integration of the effector and a distal object can be fully abolished even under conditions of full controllability.

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Susceptibility to the Rubber Hand Illusion (RHI) demonstrates that body ownership can be modulated by visuotactile inputs. In contrast to body-like images, other objects cannot be embodied suggesting that crossmodal interactions on body ownership are based on a 'goodness-of-fit' mechanism relative to one's own body. However, it is not clear whether visual self-recognition influences susceptibility to the RHI, although evidence for individual differences in the perceptual body image on the RHI suggests that this may be the case. We investigated the role of self-recognition on the subjective experience of the RHI and measured proprioceptive drift and onset time of the RHI between two groups, one with the ability to identify an image of their own hand and the other without this ability. A typical RHI response was found overall with no group difference in the subjective experience of the RHI. However, a larger proprioceptive drift and an earlier onset time for the RHI was found for the non-recognisers than the self-recognition group. Our findings provide evidence for a link between a visual representation of one's own body in long-term memory and plasticity of the body representation.

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The rubber hand illusion (RHI) is a widely applied paradigm to investigate changes in body representations. Extensive scientific interest has produced a great variability in the observed results and many contradictory findings have been reported. Taking into account the numerous variations in the experimental implementation of the RHI, many of these contradictive findings can be reconciled, but to date a thorough analysis of the methodological differences between RHI studies is lacking. Here we summarize and analyse methodological differences between RHI studies. In distinction from other reviews focusing on the integration of findings from various studies, the present paper is devoted to the differences in (i) the experimental setup, (ii) the method used to induce the RHI, (iii) the quantification of its effects, and (iv) aspects of the experimental design and data analysis. This approach will provide a reference frame for the interpretation of previous studies as well as for the design of future studies.

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Preference for use of either the left or right hand ('handedness') has been linked with modulations of perception and sensory processing-both of space and the body. Here we ask whether multisensory integration of bodily information also varies as a function of handedness. We created a spatial disparity between visual and somatosensory hand position information using the rubber hand illusion, and use the magnitude of illusory shifts in hand position (proprioceptive 'drift') as a tool to probe the weighted integration of multisensory information. First, we found drift was significantly reduced when the illusion was performed on the dominant vs. non-dominant hand. We suggest increased manual dexterity of the dominant hand causes greater representational stability and thus an increased resistance to bias by the illusion induction. Second, drift was generally greatest when the hand was in its habitual action space (i.e., near the shoulder of origin), compared to when it laterally displaced towards, or across the midline. This linear effect, however, was only significant for the dominant hand-in both left- and right-handed groups. Thus, our results reveal patterns of habitual hand action modulate drift both within a hand (drift varies with proximity to action space), and between hands (differences in drift between the dominant and non-dominant hands). In contrast, we were unable to find conclusive evidence to support, or contradict, an overall difference between left- and right-handers in susceptibility to RHI drift (i.e., total drift, collapsed across hand positions). In sum, our results provide evidence that patterns of daily activity-and the subsequent patterns of sensory input-shape multisensory integration across space.

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The rubber hand illusion (RHI) is an experimental paradigm known to produce a bodily illusion. Transcutaneous electrical nerve stimulation (TENS) combined with the RHI induces a stronger illusion than the RHI alone. Visuotactile stimulus synchrony is an important aspect of the RHI. However, the effect of TENS and visuotactile stimulus synchrony in TENS combined with the RHI remains unknown. The purpose of this study was to investigate the effects of TENS and visuotactile stimulus synchrony on the embodiment of an artificial hand when using TENS combined with the RHI. The participants underwent four experimental conditions in random order: TENS/noTENS × Synchronous/Asynchronous. TENS was set at an intensity such that it generated a feeling of electrical paresthesia in the radial nerve area of the hand but did not cause pain, i.e., 100-Hz pulse frequency, 80-µs pulse duration, and a constant pulse pattern. A visuotactile stimulus, either temporally synchronous or asynchronous, was generated using paintbrush strokes. To evaluate the outcome measures, the participants completed a questionnaire report and proprioceptive drift assessments (motor response and perceptual response). There were significant main effects of TENS and visuotactile synchrony, but no interaction between these factors, on the results of the questionnaire and the perceptual response. In contrast, there was no significant effect on the result of the motor response. These findings indicate that TENS and visuotactile synchrony might affect differently the embodiment of an artificial hand when using TENS combined with the RHI.

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In Tourette Syndrome, the expression of tics and commonly preceding premonitory sensations is associated with perturbed subjective feelings of self-control and agency. We compared responses to the Rubber Hand Illusion in 23 adults with TS and 22 controls. Both TS and control participants reported equivalent subjective embodiment of the artificial hand: feelings of ownership, location, and agency were greater during synchronous visuo-tactile stimulation, compared to asynchronous. However, individuals with TS did not manifest greater proprioceptive drift, an objective marker of embodiment observed in controls. An 'embodiment prediction error' index of the difference between subjective embodiment and objective proprioceptive drift correlated with severity of premonitory sensations. Feelings of ownership also correlated with premonitory sensation severity, and feelings of agency with tic severity. These findings suggest that subjective bodily ownership, as measured by the rubber hand illusion, contributes to susceptibility to the premonitory sensations that may be a precipitating factor in tics.

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We manipulated the sense of body ownership with the rubber hand illusion (RHI) to determine if perception of a potentially painful threat to the rubber hand can modify the mechanical pain threshold (MPT). Simultaneous tactile stimulation of the subject's concealed hand and the appropriately positioned visible rubber hand generated the illusion of false body ownership. The MPT was recorded on the left hand of the subjects before and after induction of the RHI, as well as during the phase in which the model hand was pricked with a sharp knife or touched by the blunt knife handle. The results indicate that the RHI could be successfully generated with our set-up. Mechanical stimuli were perceived as more painful in the condition where the rubber hand was simultaneously pricked with a knife. Our findings suggest that the illusion of body ownership gates nociceptive processing of potentially painful stimuli.