Proprioceptive drift in the rubber hand illusion predicts action reproduction accuracy

Proprioceptive drift in the rubber hand illusion predicts action reproduction accuracy | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Proprioceptive drift in the rubber hand illusion predicts action reproduction accuracy Masanori Sakamoto, Yuki Matsuda This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6832548/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 17 Nov, 2025 Read the published version in Experimental Brain Research → Version 1 posted You are reading this latest preprint version Abstract The neural representation of the body is highly flexible and can be altered by integrating multisensory signals in the brain. The rubber hand illusion (RHI) is a widely used paradigm to investigate this phenomenon; participants experience ownership of a rubber hand and perceive their real hand as shifting toward the rubber hand’s location, a phenomenon known as proprioceptive drift. Although individual differences in the extent of this drift are well documented, it remains unclear whether such differences are related to specific aspects of motor function. In this study, we examined the relationship between the magnitude of proprioceptive drift during the RHI and the ability of individuals to imitate and reproduce elbow movements. Our results revealed a significant correlation between the magnitude of proprioceptive drift and the accuracy of action reproduction but not imitation. These findings suggest that altered body representation may selectively influence the motor processes involved in action reproduction, highlighting the interplay between body ownership and motor control. rubber hand illusion proprioceptive drift action reproduction body representation Figures Figure 1 Figure 2 Figure 3 Introduction Neural representations of the body can be rapidly and easily altered by integrating multiple sensory signals. The rubber hand illusion (RHI) is a widely used experimental paradigm to investigate this phenomenon (Botvinick and Cohen 1998 ). In this paradigm, viewing a rubber hand being stroked in synchrony and the same direction as one’s own hidden hand induces the sensation that the rubber hand is a part of one’s body. Additionally, the perceived location of the real hand shifts toward the rubber hand, a phenomenon known as proprioceptive drift (Botvinick and Cohen 1998 ; Costantini and Haggard 2007 ; Tsakiris and Haggard 2005 ; Tsakiris et al. 2010 ). Proprioceptive drift does not occur when the stroke is asynchronous (Tsakiris and Haggard 2005 ) or in a different direction (Costantini and Haggard 2007 ). The strength of the illusion depends on the spatial distance between the rubber and real hands (Lloyd 2007 ). These findings suggest that temporal and spatial synchrony between visual and somatosensory signals is essential for illusion. When visual and tactile inputs are incongruent, visual information dominates, leading the brain to incorporate non-biological objects into its body representation (Lewis and Lloyd 2010 ; Lloyd 2007 ). Numerous studies have explored the neural mechanisms underlying the RHI, providing partial insights into the brain processes involved (Ehrsson et al. 2004 ; Ehrsson et al. 2005 ; Sakamoto and Ifuku 2021 ; Tsakiris et al. 2008 ). A few of these findings highlight the potential for the clinical application of the RHI (Hegedüs et al. 2014 ; Kammers et al. 2011 ; Kanaya et al. 2012 ; Moseley et al. 2008 ; Ramakonar et al. 2011 ). In contrast, relatively few studies have addressed individual differences in illusion strength (Abdulkarim and Ehrsson 2016 ; Asai et al. 2011 ; Marotta et al. 2016 ; Sakamoto and Ifuku 2022 ; Tsakiris et al. 2011 ). Evidence suggests that interoceptive sensitivity (Tsakiris et al. 2011 ) and empathy (Asai et al. 2011 ) are associated with variability in susceptibility to RHI. Investigating these differences offers a promising avenue to deepen our understanding of body awareness and personality traits. Whether individual differences in the RHI strength are related to specific aspects of motor skills remains unclear. Action imitation and reproduction, which are the key components of motor performance, may require individuals to generate an internal body representation and execute movements based on that representation. In such cases, flexibility in the neural representation of the body may be essential for adaptively shaping it to meet task demands. We hypothesized that the degree of malleability in body representation, as reflected by the magnitude of the RHI, is associated with an individual’s ability to imitate or reproduce actions. To test this hypothesis, we examined the relationship between proprioceptive drift during the RHI and the accuracy of imitation and reproduction of elbow movement. Methods Participants Forty healthy volunteers (10 females and 30 males) aged 20–23 years naïve to the purpose of the study participated in the experiment. All participants were right-handed, as confirmed by the Edinburgh Handedness Inventory (Oldfield 1971 ), and showed no abnormalities on physical or neurological examinations. Written informed consent was obtained from all the participants. This study was approved by the Human Research Ethics Committee of the Faculty of Education, Kumamoto University, and was conducted in accordance with the Declaration of Helsinki. Imitation Task The participants were seated comfortably in a chair with the left shoulder flexed at 90°, the left elbow fully extended (0°), and the left forearm in a supinated position. A potentiometer (TOYO PHYSICAL, Japan) was attached to the left elbow to measure the joint angles. A video monitor was positioned 100 cm in front of the participants and displayed a male model flexing his left elbow to one of ten angles (11°, 24°, 32°, 53°, 62°, 67°, 82°, 98°, 116°, or 120°) from the initial 0° position. The target angles were presented in a randomized order. The participants were instructed to observe the model’s elbow movement and then imitate the displayed angle using their left elbow. The task was repeated using the right elbow under the same conditions. Each participant completed four blocks of trials, two blocks for each arm, with each block consisting of 10 randomized trials (one for each target angle), totaling 40 trials. The inter-trial interval was 5–10 s. The block order was randomized across participants. Reproduction Task The initial posture for this task was the same as that of the imitation task. With their eyes open, participants were instructed to flex their left elbow from the fully extended position (0°) to an arbitrary angle of choice and then return to the extended position. After a delay of approximately 2–4 s, they were asked to reproduce the same elbow angle with their eyes open. Although the participants could freely select the flexion angle, they were instructed to distribute their movements roughly evenly across the 0–120° range. This procedure was repeated for the right elbow. The reproduction task consisted of four blocks of 10 trials each (two blocks per arm) for 40 trials. The order of the blocks was randomized. The interval between the trials ranged from 5 to 10 s. Rubber Hand Illusion The RHI experiment was conducted in a dimly lit room with the participants seated in a chair. The participants’ real and rubber left hands were covered with identical light blue rubber gloves to eliminate visual differences (Tsakiris and Haggard 2005 ). The participants placed their real left hand and forearm inside a wooden frame in the prone position, while the rubber left hand was positioned 19 cm medial to the real hand in the prone position. A black cloth covered the left upper arm and the area surrounding the rubber hand, allowing only the fake hand to remain visible throughout the experiment. To assess the perceived hand position (pre-position test), the participants were first asked to close their eyes while a blackboard (21 cm deep, 91 cm wide) was placed on the frame to block the view of both hands. A horizontal white line with a concealed scale (visible only to the researcher) was drawn on the board. After the board was set, the participants opened their eyes, and the experimenter moved a pointer stick laterally across the line. The participants verbally indicated when the pointer was directly above the perceived location of their left index finger. The experimenter recorded the position using a hidden scale. Following the pre-position test, the participants closed their eyes, and the board was removed. They were then instructed to open their eyes and observe the rubber hand while the experimenter applied synchronous or asynchronous tactile stimulation to the real and rubber hands for 3 min using two identical paintbrushes. In the congruent condition, the strokes were temporally and spatially matched. In the incongruent condition, the strokes were mismatched in terms of timing and/or location. Each condition was repeated three times in a randomized order with a 5-min rest period between sessions. After each stimulation session, the perceived position of the left index finger was assessed using the same procedure (post-position test). Following the final post-position test, participants completed a questionnaire based on Botvinick and Cohen’s ( 1998 ) original RHI study. The questionnaire included the following eight statements. S1: It seemed as if I were feeling the touch of the paintbrush in the location where I saw the rubber hand touched. S2: It seemed as though the touch I felt was caused by the paintbrush touching the rubber hand. S3: I felt as if the rubber hand were my hand. S4: It felt as if my (real) hand were drifting toward the rubber hand. S5: It seemed as if I might have more than one left/right hand or arm. S6: It seemed as if the touch I was feeling came from somewhere between my own hand and the rubber hand. S7: It felt as if my (real) hand were turning 'rubbery.' S8: It appeared (visually) as if the rubber hand were drifting toward my hand. Participants rated each statement on a 10-point scale, ranging from 1 (strongly disagree) to 10 (strongly agree). Data Analysis and Statistics For the imitation task, the position errors were calculated by subtracting the model’s elbow angle (as shown in the video) from the participant’s elbow angle. Errors were averaged for each block. For the reproduction task, the position errors were calculated by subtracting the angle of the first elbow flexion from that of the second (reproduced) flexion, and the values were averaged for each block. Both absolute errors (magnitude of the difference) and constant errors (directional bias) were computed. To assess the modulation of these errors, a two-way repeated measures analysis of variance (ANOVA) was performed with two factors: task type (imitation vs. reproduction) and hand used (left vs. right). To analyze the questionnaire data, the Friedman test was used to assess the effect of the stroking condition (congruent vs. incongruent) on each questionnaire item. When significant effects were observed, post-hoc comparisons were conducted using the Wilcoxon signed-rank test. Proprioceptive drift, defined as a shift in the perceived hand position toward the rubber hand, was calculated for each stroking condition (congruent and incongruent) by subtracting the pre-position from the post-position judgment. The drift was measured three times per condition, and the values were averaged. As in previous studies (e.g., Botvinick and Cohen 1998 ; Tsakiris and Haggard 2005 ), this average drift was used as an objective measure of illusion strength. A paired t-test was conducted to compare the magnitude of proprioceptive drift between the two stroking conditions. Additionally, we computed the “net” magnitude of the RHI by subtracting the incongruent values from the congruent values for both proprioceptive drift and questionnaire ratings (following Sakamoto and Ifuku 2022 ; Sakamoto et al. 2024 ). To explore the relationship between RHI and motor performance, linear regression analyses were performed between net proprioceptive drift and task performance (absolute and constant errors in the imitation and reproduction tasks). Pearson’s correlation coefficient ( r ) were calculated. The relationships between net questionnaire ratings and task performance were assessed using Spearman’s rank correlation coefficients ( ρ ). Statistical significance was set at p < 0.05. All analyses were performed using the IBM SPSS Statistics software. Results Figure 1 A shows the individual and mean absolute errors of the imitation and reproduction tasks. A two-way repeated-measures ANOVA revealed a significant main effect of task type ( F (1, 39) = 106.5, p < 0.001, η ² = 0.73), indicating that absolute errors differed significantly between tasks. However, no significant main effect of hand used ( F (1, 39) = 0.37, p = 0.55, η ² = 0.01), and no significant interaction between task and hand ( F (1, 39) = 0.07, p = 0.80, η ² = 0.002) were observed. Figure 1 B shows the individual and mean constant errors. The two-way ANOVA revealed a significant main effect of task type ( F (1, 39) = 51.9, p < 0.001, η ² = 0.57). No significant main effect of hand ( F (1, 39) = 0.72, p = 0.40, η ² = 0.02), and no significant interaction between task and hand ( F (1, 39) = 0.0003, p = 0.99, η ² = 0.00007) were observed. Figure 2 A shows the magnitude of proprioceptive drift. A paired t -test showed that drift was significantly greater following congruent stroking than incongruent stroking ( p < 0.001), indicating a stronger illusion under congruent conditions. Figure 2 B shows the ratings obtained from the RHI questionnaire. The Friedman test revealed significant modulation of ratings under the congruent condition ( χ ² = 158.1, p < 0.001). Bonferroni-corrected pairwise comparisons showed that the ratings for statements 1, 2, and 3 were significantly higher than those for all other items ( p < 0.001 for all). In contrast, the Friedman test for the incongruent condition showed no significant modulation across items ( χ ² = 10.3, p = 0.17). Wilcoxon signed-rank tests revealed that the ratings for statements 1, 2, and 3 were significantly higher during congruent stroking than incongruent stroking ( p < 0.001 for all). Figure 3 shows the relationship between the net magnitude of proprioceptive drift and absolute errors in the imitation and reproduction tasks. A significant negative correlation was observed only between drift and absolute error in the reproduction task with the left hand ( r = − 0.46, p = 0.003; Figure. 3D). No significant correlations were observed in the imitation with the right hand ( r = 0.11, p = 0.48; Fig. 3 A), imitation with the left hand (Fig. 3 B), or reproduction with the right hand (Fig. 3 C). No significant correlations were observed between the proprioceptive drift and constant errors in any of the four conditions (data not shown). Additionally, no significant correlations were observed between the RHI questionnaire ratings and absolute or constant errors in the imitation or reproduction tasks (data not shown). Discussion In this study, we investigated the relationship between changes in the neural representations of the body and the ability to reproduce or imitate actions. We observed a significant negative correlation between the strength of proprioceptive drift during the RHI and angular errors in the reproduction task. In contrast, no such correlation was observed in the imitation task. Additionally, the subjective ratings from the RHI questionnaire did not correlate with the accuracy of either task. These findings suggest that individuals who exhibit greater susceptibility to changes in body representation may possess superior action-reproduction abilities. Thus, modulation of the neural representation of the body may be a key factor in action reproduction. As shown in Fig. 2 A, the magnitude of the proprioceptive drift was significantly larger in the congruent stroking condition than in the incongruent condition. Furthermore, questionnaire items 1, 2, and 3, which are commonly associated with changes in the sense of body ownership, were rated higher than other items in the congruent condition (Fig. 2 B). These findings confirm that the RHI was successfully induced in the current study, which aligns with previous studies (Botvinick and Cohen 1998 ; Tsakiris and Haggard 2005 ; Tsakiris et al. 2010 ). A significant correlation between proprioceptive drift and reproduction accuracy was observed only in the left-hand reproduction task, where the same limb was involved in the RHI (Fig. 3 D). This suggests that RHI-induced modulation of motor performance is limb-specific and cannot be generalized to limbs that are not directly involved in the illusion. Moreover, a significant correlation was observed for absolute but not constant errors. This implies that the strength of the illusion is related to the overall accuracy of reproducing a movement rather than a directional bias (i.e., whether the reproduced angle was too flexed or extended). Although the underlying neural mechanisms could not be directly identified in this study, previous research has suggested that the posterior parietal cortex may be involved. Activity in this region has been associated with artificial limb ownership during the RHI (Ehrsson et al. 2004 ; Tsakiris et al. 2008 ). The parietal cortex has been implicated in various spatial functions, including 3D object recognition (Durand et al. 2009 ), postural updating (Parkinson et al. 2010 ), and spatial navigation (Calton and Taube 2009 ). As the reproduction task required spatial encoding of elbow angles, it is plausible that individuals exhibiting greater proprioceptive drift, reflecting greater modulation of spatial body representation, were better able to reproduce the task accurately. The difference in the findings between the reproduction and imitation tasks may be attributed to the nature of the sensory information involved. In the reproduction task, participants used visual and somatosensory inputs during the first flexion to construct a reference body image, which was then used to guide the second flexion. This process involves the integration of multiple sensory modalities, a key requirement for inducing RHI (Tsakiris 2010 ), and likely reflects dynamic updates to one’s body representation. In contrast, the imitation task relies only on visual information obtained by observing another person’s actions. This lack of somatosensory integration may explain why no correlation was observed between the RHI strength and imitation performance. The subjective ratings of body ownership in the RHI questionnaire did not significantly correlate with the performance on the reproduction task. While both proprioceptive drift and ownership ratings are commonly used indices of RHI strength (Botvinick and Cohen 1998 ), previous studies have reported a dissociation between these measures (e.g., Abdulkarim and Ehrsson 2016 ; Holle et al. 2011 ; Rohde et al. 2011 ). Rohde et al. ( 2011 ) proposed that proprioceptive drift reflects a spatial updating process driven by synchronous visuotactile inputs that are distinct from the conscious feeling of limb ownership. Our findings support this view, suggesting that the spatial updating of body position, rather than subjective ownership, is more closely related to the ability to accurately reproduce actions. Previous studies on the RHI have paid limited attention to individual differences in the magnitude of the illusion and have rarely explored how these differences relate to motor abilities. This study demonstrated that changes in the neural representation of the body, as reflected by the magnitude of proprioceptive drift, are associated with the ability to accurately reproduce actions. These findings suggest that individual variability in body representation plasticity may be a critical factor in motor reproductive performance. Moreover, our results highlighted the potential utility of the RHI paradigm as an objective tool for assessing specific aspects of motor function related to body representation. Declarations Conflict of interest The authors declare no competing interests. Funding This work was supported by JSPS KAKENHI Grant Number JP24700610. Author Contribution M.S. and Y.M. designed the experiment; Y.M. performed data collection; M.S. and Y.M. analyzed data; M.S. prepared all figures; M.S. wrote the article. All authors reviewed the manuscript. Acknowledgement This work was supported by JSPS KAKENHI Grant Number JP24700610. The authors would like to thank Editage (www.editage.jp) for English language editing. Data Availability Data will be made available on request. References Abdulkarim Z, Ehrsson HH (2016) No causal link between changes in hand position sense and feeling of limb ownership in the rubber hand illusion. Atten Percept Psychophys 78:707–720. https://doi.org/10.3758/s13414-015-1016-0 Asai T, Mao Z, Sugimori E, Tanno Y (2011) Rubber hand illusion, empathy, and schizotypal experiences in terms of self-other representations. Conscious Cogn 20:1744–17850. https://doi.org/10.1016/j.concog.2011.02.005 Botvinick M, Cohen J (1998) Rubber hands 'feel' touch that eyes see. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6832548","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":470359108,"identity":"5491977a-0542-4dcd-9e0c-9fb50b09b7a1","order_by":0,"name":"Masanori Sakamoto","email":"data:image/png;base64,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","orcid":"","institution":"Kumamoto University","correspondingAuthor":true,"prefix":"","firstName":"Masanori","middleName":"","lastName":"Sakamoto","suffix":""},{"id":470359110,"identity":"896d824f-9cc3-4dc1-8709-364212ca9fda","order_by":1,"name":"Yuki Matsuda","email":"","orcid":"","institution":"Kumamoto University","correspondingAuthor":false,"prefix":"","firstName":"Yuki","middleName":"","lastName":"Matsuda","suffix":""}],"badges":[],"createdAt":"2025-06-06 01:23:14","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6832548/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6832548/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s00221-025-07192-8","type":"published","date":"2025-11-17T15:57:30+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":84684445,"identity":"a25c74d7-df40-465b-8637-10fcca1b6d97","added_by":"auto","created_at":"2025-06-16 08:46:03","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":73529,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eAbsolute (A) and constant (B) errors in the imitation and reproduction tasks\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eDots represent individual data points, and superimposed bar graphs indicate group means. In panel (B), the positive and negative constant error values in the imitation task reflect that the participants’ elbow angles were more flexed or extended, respectively, than the model’s angle. For the reproduction task, positive and negative constant error values indicated that the second elbow flexion was more extended than the first flexion.\u003c/p\u003e","description":"","filename":"Figure11.png","url":"https://assets-eu.researchsquare.com/files/rs-6832548/v1/97f259c4a6408784957088f4.png"},{"id":84684446,"identity":"1cae20f3-609b-42c3-b065-6deeffa43145","added_by":"auto","created_at":"2025-06-16 08:46:03","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":37870,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003e(A) Proprioceptive drift and (B) questionnaire ratings in the rubber hand illusion\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e(A) Group mean and standard deviation of proprioceptive drift, measured in centimeters. Positive values indicate drift toward the rubber hand.\u003c/p\u003e\n\u003cp\u003e(B) Mean ratings (± standard deviation) of the RHI questionnaire items under congruent and incongruent strokingconditions. Participants rated their agreement with each statement on a 10-point scale (1 = strongly disagree, 10 = strongly agree).\u003c/p\u003e","description":"","filename":"Figure21.png","url":"https://assets-eu.researchsquare.com/files/rs-6832548/v1/fb8bb8530e4e39a6871839d3.png"},{"id":84682595,"identity":"8220821f-c160-4d1c-bf2f-80c902886059","added_by":"auto","created_at":"2025-06-16 08:38:03","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":53085,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eRelationship between the “net” magnitude of proprioceptive drift and absolute errors in the imitation (A, B) and reproduction (C, D) tasks\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe abscissa represents the “net” proprioceptive drift, defined as the difference in drift magnitude between congruent and incongruent strokingconditions. Positive values indicated greater drift during congruent stroking. The correlation analyses were performed separately for each task and limb.\u003c/p\u003e","description":"","filename":"Figure31.png","url":"https://assets-eu.researchsquare.com/files/rs-6832548/v1/1db5a199ad3fbd3d41c807f2.png"},{"id":96650091,"identity":"949b39ab-308d-4a33-9ae4-1d8663505833","added_by":"auto","created_at":"2025-11-24 16:07:07","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":654915,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6832548/v1/86c0e912-df26-4712-9bb5-50d289bfe978.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Proprioceptive drift in the rubber hand illusion predicts action reproduction accuracy","fulltext":[{"header":"Introduction","content":"\u003cp\u003eNeural representations of the body can be rapidly and easily altered by integrating multiple sensory signals. The rubber hand illusion (RHI) is a widely used experimental paradigm to investigate this phenomenon (Botvinick and Cohen \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). In this paradigm, viewing a rubber hand being stroked in synchrony and the same direction as one\u0026rsquo;s own hidden hand induces the sensation that the rubber hand is a part of one\u0026rsquo;s body. Additionally, the perceived location of the real hand shifts toward the rubber hand, a phenomenon known as proprioceptive drift (Botvinick and Cohen \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Costantini and Haggard \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Tsakiris and Haggard \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Tsakiris et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Proprioceptive drift does not occur when the stroke is asynchronous (Tsakiris and Haggard \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2005\u003c/span\u003e) or in a different direction (Costantini and Haggard \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). The strength of the illusion depends on the spatial distance between the rubber and real hands (Lloyd \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). These findings suggest that temporal and spatial synchrony between visual and somatosensory signals is essential for illusion. When visual and tactile inputs are incongruent, visual information dominates, leading the brain to incorporate non-biological objects into its body representation (Lewis and Lloyd \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Lloyd \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eNumerous studies have explored the neural mechanisms underlying the RHI, providing partial insights into the brain processes involved (Ehrsson et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Ehrsson et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Sakamoto and Ifuku \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Tsakiris et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). A few of these findings highlight the potential for the clinical application of the RHI (Heged\u0026uuml;s et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Kammers et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Kanaya et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Moseley et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Ramakonar et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). In contrast, relatively few studies have addressed individual differences in illusion strength (Abdulkarim and Ehrsson \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Asai et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Marotta et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Sakamoto and Ifuku \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Tsakiris et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Evidence suggests that interoceptive sensitivity (Tsakiris et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) and empathy (Asai et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) are associated with variability in susceptibility to RHI. Investigating these differences offers a promising avenue to deepen our understanding of body awareness and personality traits.\u003c/p\u003e \u003cp\u003eWhether individual differences in the RHI strength are related to specific aspects of motor skills remains unclear. Action imitation and reproduction, which are the key components of motor performance, may require individuals to generate an internal body representation and execute movements based on that representation. In such cases, flexibility in the neural representation of the body may be essential for adaptively shaping it to meet task demands. We hypothesized that the degree of malleability in body representation, as reflected by the magnitude of the RHI, is associated with an individual\u0026rsquo;s ability to imitate or reproduce actions. To test this hypothesis, we examined the relationship between proprioceptive drift during the RHI and the accuracy of imitation and reproduction of elbow movement.\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eParticipants\u003c/h2\u003e \u003cp\u003eForty healthy volunteers (10 females and 30 males) aged 20\u0026ndash;23 years na\u0026iuml;ve to the purpose of the study participated in the experiment. All participants were right-handed, as confirmed by the Edinburgh Handedness Inventory (Oldfield \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1971\u003c/span\u003e), and showed no abnormalities on physical or neurological examinations. Written informed consent was obtained from all the participants. This study was approved by the Human Research Ethics Committee of the Faculty of Education, Kumamoto University, and was conducted in accordance with the Declaration of Helsinki.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eImitation Task\u003c/h3\u003e\n\u003cp\u003eThe participants were seated comfortably in a chair with the left shoulder flexed at 90\u0026deg;, the left elbow fully extended (0\u0026deg;), and the left forearm in a supinated position. A potentiometer (TOYO PHYSICAL, Japan) was attached to the left elbow to measure the joint angles. A video monitor was positioned 100 cm in front of the participants and displayed a male model flexing his left elbow to one of ten angles (11\u0026deg;, 24\u0026deg;, 32\u0026deg;, 53\u0026deg;, 62\u0026deg;, 67\u0026deg;, 82\u0026deg;, 98\u0026deg;, 116\u0026deg;, or 120\u0026deg;) from the initial 0\u0026deg; position. The target angles were presented in a randomized order.\u003c/p\u003e \u003cp\u003eThe participants were instructed to observe the model\u0026rsquo;s elbow movement and then imitate the displayed angle using their left elbow. The task was repeated using the right elbow under the same conditions. Each participant completed four blocks of trials, two blocks for each arm, with each block consisting of 10 randomized trials (one for each target angle), totaling 40 trials. The inter-trial interval was 5\u0026ndash;10 s. The block order was randomized across participants.\u003c/p\u003e\n\u003ch3\u003eReproduction Task\u003c/h3\u003e\n\u003cp\u003eThe initial posture for this task was the same as that of the imitation task. With their eyes open, participants were instructed to flex their left elbow from the fully extended position (0\u0026deg;) to an arbitrary angle of choice and then return to the extended position. After a delay of approximately 2\u0026ndash;4 s, they were asked to reproduce the same elbow angle with their eyes open. Although the participants could freely select the flexion angle, they were instructed to distribute their movements roughly evenly across the 0\u0026ndash;120\u0026deg; range. This procedure was repeated for the right elbow.\u003c/p\u003e \u003cp\u003eThe reproduction task consisted of four blocks of 10 trials each (two blocks per arm) for 40 trials. The order of the blocks was randomized. The interval between the trials ranged from 5 to 10 s.\u003c/p\u003e\n\u003ch3\u003eRubber Hand Illusion\u003c/h3\u003e\n\u003cp\u003eThe RHI experiment was conducted in a dimly lit room with the participants seated in a chair. The participants\u0026rsquo; real and rubber left hands were covered with identical light blue rubber gloves to eliminate visual differences (Tsakiris and Haggard \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). The participants placed their real left hand and forearm inside a wooden frame in the prone position, while the rubber left hand was positioned 19 cm medial to the real hand in the prone position. A black cloth covered the left upper arm and the area surrounding the rubber hand, allowing only the fake hand to remain visible throughout the experiment.\u003c/p\u003e \u003cp\u003eTo assess the perceived hand position (pre-position test), the participants were first asked to close their eyes while a blackboard (21 cm deep, 91 cm wide) was placed on the frame to block the view of both hands. A horizontal white line with a concealed scale (visible only to the researcher) was drawn on the board. After the board was set, the participants opened their eyes, and the experimenter moved a pointer stick laterally across the line. The participants verbally indicated when the pointer was directly above the perceived location of their left index finger. The experimenter recorded the position using a hidden scale.\u003c/p\u003e \u003cp\u003eFollowing the pre-position test, the participants closed their eyes, and the board was removed. They were then instructed to open their eyes and observe the rubber hand while the experimenter applied synchronous or asynchronous tactile stimulation to the real and rubber hands for 3 min using two identical paintbrushes. In the congruent condition, the strokes were temporally and spatially matched. In the incongruent condition, the strokes were mismatched in terms of timing and/or location. Each condition was repeated three times in a randomized order with a 5-min rest period between sessions. After each stimulation session, the perceived position of the left index finger was assessed using the same procedure (post-position test).\u003c/p\u003e \u003cp\u003eFollowing the final post-position test, participants completed a questionnaire based on Botvinick and Cohen\u0026rsquo;s (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1998\u003c/span\u003e) original RHI study. The questionnaire included the following eight statements.\u003c/p\u003e \u003cp\u003eS1: It seemed as if I were feeling the touch of the paintbrush in the location where I saw the rubber hand touched.\u003c/p\u003e \u003cp\u003eS2: It seemed as though the touch I felt was caused by the paintbrush touching the rubber hand.\u003c/p\u003e \u003cp\u003eS3: I felt as if the rubber hand were my hand.\u003c/p\u003e \u003cp\u003eS4: It felt as if my (real) hand were drifting toward the rubber hand.\u003c/p\u003e \u003cp\u003eS5: It seemed as if I might have more than one left/right hand or arm.\u003c/p\u003e \u003cp\u003eS6: It seemed as if the touch I was feeling came from somewhere between my own hand and the rubber hand.\u003c/p\u003e \u003cp\u003eS7: It felt as if my (real) hand were turning 'rubbery.'\u003c/p\u003e \u003cp\u003eS8: It appeared (visually) as if the rubber hand were drifting toward my hand.\u003c/p\u003e \u003cp\u003eParticipants rated each statement on a 10-point scale, ranging from 1 (strongly disagree) to 10 (strongly agree).\u003c/p\u003e\n\u003ch3\u003eData Analysis and Statistics\u003c/h3\u003e\n\u003cp\u003eFor the imitation task, the position errors were calculated by subtracting the model\u0026rsquo;s elbow angle (as shown in the video) from the participant\u0026rsquo;s elbow angle. Errors were averaged for each block. For the reproduction task, the position errors were calculated by subtracting the angle of the first elbow flexion from that of the second (reproduced) flexion, and the values were averaged for each block.\u003c/p\u003e \u003cp\u003eBoth absolute errors (magnitude of the difference) and constant errors (directional bias) were computed. To assess the modulation of these errors, a two-way repeated measures analysis of variance (ANOVA) was performed with two factors: task type (imitation vs. reproduction) and hand used (left vs. right).\u003c/p\u003e \u003cp\u003eTo analyze the questionnaire data, the Friedman test was used to assess the effect of the stroking condition (congruent vs. incongruent) on each questionnaire item. When significant effects were observed, post-hoc comparisons were conducted using the Wilcoxon signed-rank test.\u003c/p\u003e \u003cp\u003eProprioceptive drift, defined as a shift in the perceived hand position toward the rubber hand, was calculated for each stroking condition (congruent and incongruent) by subtracting the pre-position from the post-position judgment. The drift was measured three times per condition, and the values were averaged. As in previous studies (e.g., Botvinick and Cohen \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Tsakiris and Haggard \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), this average drift was used as an objective measure of illusion strength.\u003c/p\u003e \u003cp\u003eA paired t-test was conducted to compare the magnitude of proprioceptive drift between the two stroking conditions. Additionally, we computed the \u0026ldquo;net\u0026rdquo; magnitude of the RHI by subtracting the incongruent values from the congruent values for both proprioceptive drift and questionnaire ratings (following Sakamoto and Ifuku \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Sakamoto et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo explore the relationship between RHI and motor performance, linear regression analyses were performed between net proprioceptive drift and task performance (absolute and constant errors in the imitation and reproduction tasks). Pearson\u0026rsquo;s correlation coefficient (\u003cem\u003er\u003c/em\u003e) were calculated. The relationships between net questionnaire ratings and task performance were assessed using Spearman\u0026rsquo;s rank correlation coefficients (\u003cem\u003eρ\u003c/em\u003e).\u003c/p\u003e \u003cp\u003eStatistical significance was set at \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05. All analyses were performed using the IBM SPSS Statistics software.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eFigure \u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA shows the individual and mean absolute errors of the imitation and reproduction tasks. A two-way repeated-measures ANOVA revealed a significant main effect of task type (\u003cem\u003eF\u003c/em\u003e(1, 39)\u0026thinsp;=\u0026thinsp;106.5, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001, \u003cem\u003eη\u003c/em\u003e\u0026sup2; = 0.73), indicating that absolute errors differed significantly between tasks. However, no significant main effect of hand used (\u003cem\u003eF\u003c/em\u003e(1, 39)\u0026thinsp;=\u0026thinsp;0.37, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.55, \u003cem\u003eη\u003c/em\u003e\u0026sup2; = 0.01), and no significant interaction between task and hand (\u003cem\u003eF\u003c/em\u003e(1, 39)\u0026thinsp;=\u0026thinsp;0.07, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.80, \u003cem\u003eη\u003c/em\u003e\u0026sup2; = 0.002) were observed.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eFigure \u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB shows the individual and mean constant errors. The two-way ANOVA revealed a significant main effect of task type (\u003cem\u003eF\u003c/em\u003e(1, 39)\u0026thinsp;=\u0026thinsp;51.9, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001, \u003cem\u003eη\u003c/em\u003e\u0026sup2; = 0.57). No significant main effect of hand (\u003cem\u003eF\u003c/em\u003e(1, 39)\u0026thinsp;=\u0026thinsp;0.72, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.40, \u003cem\u003eη\u003c/em\u003e\u0026sup2; = 0.02), and no significant interaction between task and hand (\u003cem\u003eF\u003c/em\u003e(1, 39)\u0026thinsp;=\u0026thinsp;0.0003, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.99, \u003cem\u003eη\u003c/em\u003e\u0026sup2; = 0.00007) were observed.\u003c/p\u003e \u003cp\u003eFigure \u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA shows the magnitude of proprioceptive drift. A paired \u003cem\u003et\u003c/em\u003e-test showed that drift was significantly greater following congruent stroking than incongruent stroking (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), indicating a stronger illusion under congruent conditions.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eFigure \u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eB shows the ratings obtained from the RHI questionnaire. The Friedman test revealed significant modulation of ratings under the congruent condition (\u003cem\u003eχ\u003c/em\u003e\u0026sup2; = 158.1, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Bonferroni-corrected pairwise comparisons showed that the ratings for statements 1, 2, and 3 were significantly higher than those for all other items (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001 for all). In contrast, the Friedman test for the incongruent condition showed no significant modulation across items (\u003cem\u003eχ\u003c/em\u003e\u0026sup2; = 10.3, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.17). Wilcoxon signed-rank tests revealed that the ratings for statements 1, 2, and 3 were significantly higher during congruent stroking than incongruent stroking (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001 for all).\u003c/p\u003e \u003cp\u003eFigure \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e shows the relationship between the net magnitude of proprioceptive drift and absolute errors in the imitation and reproduction tasks. A significant negative correlation was observed only between drift and absolute error in the reproduction task with the left hand (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;\u0026minus;\u0026thinsp;0.46, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.003; Figure. 3D). No significant correlations were observed in the imitation with the right hand (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.11, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.48; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eA), imitation with the left hand (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eB), or reproduction with the right hand (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eC).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eNo significant correlations were observed between the proprioceptive drift and constant errors in any of the four conditions (data not shown). Additionally, no significant correlations were observed between the RHI questionnaire ratings and absolute or constant errors in the imitation or reproduction tasks (data not shown).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn this study, we investigated the relationship between changes in the neural representations of the body and the ability to reproduce or imitate actions. We observed a significant negative correlation between the strength of proprioceptive drift during the RHI and angular errors in the reproduction task. In contrast, no such correlation was observed in the imitation task. Additionally, the subjective ratings from the RHI questionnaire did not correlate with the accuracy of either task. These findings suggest that individuals who exhibit greater susceptibility to changes in body representation may possess superior action-reproduction abilities. Thus, modulation of the neural representation of the body may be a key factor in action reproduction.\u003c/p\u003e \u003cp\u003eAs shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA, the magnitude of the proprioceptive drift was significantly larger in the congruent stroking condition than in the incongruent condition. Furthermore, questionnaire items 1, 2, and 3, which are commonly associated with changes in the sense of body ownership, were rated higher than other items in the congruent condition (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eB). These findings confirm that the RHI was successfully induced in the current study, which aligns with previous studies (Botvinick and Cohen \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Tsakiris and Haggard \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Tsakiris et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2010\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eA significant correlation between proprioceptive drift and reproduction accuracy was observed only in the left-hand reproduction task, where the same limb was involved in the RHI (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eD). This suggests that RHI-induced modulation of motor performance is limb-specific and cannot be generalized to limbs that are not directly involved in the illusion. Moreover, a significant correlation was observed for absolute but not constant errors. This implies that the strength of the illusion is related to the overall accuracy of reproducing a movement rather than a directional bias (i.e., whether the reproduced angle was too flexed or extended).\u003c/p\u003e \u003cp\u003eAlthough the underlying neural mechanisms could not be directly identified in this study, previous research has suggested that the posterior parietal cortex may be involved. Activity in this region has been associated with artificial limb ownership during the RHI (Ehrsson et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Tsakiris et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). The parietal cortex has been implicated in various spatial functions, including 3D object recognition (Durand et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), postural updating (Parkinson et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), and spatial navigation (Calton and Taube \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). As the reproduction task required spatial encoding of elbow angles, it is plausible that individuals exhibiting greater proprioceptive drift, reflecting greater modulation of spatial body representation, were better able to reproduce the task accurately.\u003c/p\u003e \u003cp\u003eThe difference in the findings between the reproduction and imitation tasks may be attributed to the nature of the sensory information involved. In the reproduction task, participants used visual and somatosensory inputs during the first flexion to construct a reference body image, which was then used to guide the second flexion. This process involves the integration of multiple sensory modalities, a key requirement for inducing RHI (Tsakiris \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), and likely reflects dynamic updates to one\u0026rsquo;s body representation. In contrast, the imitation task relies only on visual information obtained by observing another person\u0026rsquo;s actions. This lack of somatosensory integration may explain why no correlation was observed between the RHI strength and imitation performance.\u003c/p\u003e \u003cp\u003eThe subjective ratings of body ownership in the RHI questionnaire did not significantly correlate with the performance on the reproduction task. While both proprioceptive drift and ownership ratings are commonly used indices of RHI strength (Botvinick and Cohen \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1998\u003c/span\u003e), previous studies have reported a dissociation between these measures (e.g., Abdulkarim and Ehrsson \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Holle et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Rohde et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Rohde et al. (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) proposed that proprioceptive drift reflects a spatial updating process driven by synchronous visuotactile inputs that are distinct from the conscious feeling of limb ownership. Our findings support this view, suggesting that the spatial updating of body position, rather than subjective ownership, is more closely related to the ability to accurately reproduce actions.\u003c/p\u003e \u003cp\u003ePrevious studies on the RHI have paid limited attention to individual differences in the magnitude of the illusion and have rarely explored how these differences relate to motor abilities. This study demonstrated that changes in the neural representation of the body, as reflected by the magnitude of proprioceptive drift, are associated with the ability to accurately reproduce actions. These findings suggest that individual variability in body representation plasticity may be a critical factor in motor reproductive performance. Moreover, our results highlighted the potential utility of the RHI paradigm as an objective tool for assessing specific aspects of motor function related to body representation.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eConflict of interest\u003c/h2\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e\n\u003ch2\u003eFunding\u003c/h2\u003e\n\u003cp\u003eThis work was supported by JSPS KAKENHI Grant Number JP24700610.\u003c/p\u003e\n\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\n\u003cp\u003eM.S. and Y.M. designed the experiment; Y.M. performed data collection; M.S. and Y.M. analyzed data; M.S. prepared all figures; M.S. wrote the article. All authors reviewed the manuscript.\u003c/p\u003e\n\u003ch2\u003eAcknowledgement\u003c/h2\u003e\n\u003cp\u003eThis work was supported by JSPS KAKENHI Grant Number JP24700610. The authors would like to thank Editage (www.editage.jp) for English language editing.\u003c/p\u003e\n\u003ch2\u003eData Availability\u003c/h2\u003e\n\u003cp\u003eData will be made available on request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAbdulkarim Z, Ehrsson HH (2016) No causal link between changes in hand position sense and feeling of limb ownership in the rubber hand illusion. Atten Percept Psychophys 78:707\u0026ndash;720. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3758/s13414-015-1016-0\u003c/span\u003e\u003cspan address=\"10.3758/s13414-015-1016-0\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAsai T, Mao Z, Sugimori E, Tanno Y (2011) Rubber hand illusion, empathy, and schizotypal experiences in terms of self-other representations. 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Proc Biol Sci 278:2470\u0026ndash;2476. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1098/rspb.2010.2547\u003c/span\u003e\u003cspan address=\"10.1098/rspb.2010.2547\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"rubber hand illusion, proprioceptive drift, action reproduction, body representation","lastPublishedDoi":"10.21203/rs.3.rs-6832548/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6832548/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe neural representation of the body is highly flexible and can be altered by integrating multisensory signals in the brain. The rubber hand illusion (RHI) is a widely used paradigm to investigate this phenomenon; participants experience ownership of a rubber hand and perceive their real hand as shifting toward the rubber hand\u0026rsquo;s location, a phenomenon known as proprioceptive drift. Although individual differences in the extent of this drift are well documented, it remains unclear whether such differences are related to specific aspects of motor function. In this study, we examined the relationship between the magnitude of proprioceptive drift during the RHI and the ability of individuals to imitate and reproduce elbow movements. Our results revealed a significant correlation between the magnitude of proprioceptive drift and the accuracy of action reproduction but not imitation. These findings suggest that altered body representation may selectively influence the motor processes involved in action reproduction, highlighting the interplay between body ownership and motor control.\u003c/p\u003e","manuscriptTitle":"Proprioceptive drift in the rubber hand illusion predicts action reproduction accuracy","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-16 08:37:59","doi":"10.21203/rs.3.rs-6832548/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"325e5172-30e7-48be-911e-591b16562d02","owner":[],"postedDate":"June 16th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2025-11-24T16:00:55+00:00","versionOfRecord":{"articleIdentity":"rs-6832548","link":"https://doi.org/10.1007/s00221-025-07192-8","journal":{"identity":"experimental-brain-research","isVorOnly":false,"title":"Experimental Brain Research"},"publishedOn":"2025-11-17 15:57:30","publishedOnDateReadable":"November 17th, 2025"},"versionCreatedAt":"2025-06-16 08:37:59","video":"","vorDoi":"10.1007/s00221-025-07192-8","vorDoiUrl":"https://doi.org/10.1007/s00221-025-07192-8","workflowStages":[]},"version":"v1","identity":"rs-6832548","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6832548","identity":"rs-6832548","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}