Referral of Touch and Ownership between the Hands and the Role of the Somatosensory Cortices

Recent studies have shown that the feeling of body ownership can be fooled by simple visuo-tactile manipulations. Perceptual illusions have been reported in which participants sense phantom touch seen on a rubber hand (rubber hand illusion). While previous studies used homologous limbs for those experiments, we here examined an illusion where people feel phantom touch on a left rubber hand when they see it brushed simultaneously with brushes applied to their right hand. Thus, we investigated a referral of touch from the right to the left hand (across the body midline). Since it is known from animal studies that tactile illusions may alter early sensory processing, we expected a modulation of the primary somatosensory cortex (SI) corresponding to this illusion. Neuromagnetic source imaging of the functional topographic organization in SI showed a shift in left SI, associated with the strength of the referral of touch. Hence, we argue that SI seems to be closely associated with this perceptual illusion. The results suggest that the transfer of tactile information across the body midline could be mediated by neurons with bilateral tactile receptive fields (most likely BA2).     

Rubber hands feel touch, but not in blind individuals

Psychology and neuroscience have a long-standing tradition of studying blind individuals to investigate how visual experience shapes perception of the external world. Here, we study how blind people experience their own body by exposing them to a multisensory body illusion: the somatic rubber hand illusion. In this illusion, healthy blindfolded participants experience that they are touching their own right hand with their left index finger, when in fact they are touching a rubber hand with their left index finger while the experimenter touches their right hand in a synchronized manner (Ehrsson et al. 2005). We compared the strength of this illusion in a group of blind individuals (n = 10), all of whom had experienced severe visual impairment or complete blindness from birth, and a group of age-matched blindfolded sighted participants (n = 12). The illusion was quantified subjectively using questionnaires and behaviorally by asking participants to point to the felt location of the right hand. The results showed that the sighted participants experienced a strong illusion, whereas the blind participants experienced no illusion at all, a difference that was evident in both tests employed. A further experiment testing the participants' basic ability to localize the right hand in space without vision (proprioception) revealed no difference between the two groups. Taken together, these results suggest that blind individuals with impaired visual development have a more veridical percept of self-touch and a less flexible and dynamic representation of their own body in space compared to sighted individuals. We speculate that the multisensory brain systems that re-map somatosensory signals onto external reference frames are less developed in blind individuals and therefore do not allow efficient fusion of tactile and proprioceptive signals from the two upper limbs into a single illusory experience of self-touch as in sighted individuals.     

The body beyond the body: expectation of a sensory event is enough to induce ownership over a fake hand

More than 100 papers have been published on the rubber hand illusion since its discovery 14 years ago. The illusion has been proposed as a demonstration that the body is distinguished from other objects by its participation in specific forms of intermodal perceptual correlation. Here, we radically challenge this view by claiming that perceptual correlation is not necessary to produce the experience of this body as mine. Each of 15 participants was seated with his/her right arm resting upon a table just below another smaller table. Thus, the real hand was hidden from the participant''s view and a life-sized rubber model of a right hand was placed on the small table in front of the participant. The participant observed the experimenter''s hand while approaching—without touching—the rubber hand. Phenomenology of the illusion was measured by means of skin conductance response and questionnaire. Both measures indicated that participants experienced the illusion that the experimenter''s hand was about to touch their hidden hand rather than the rubber hand, as if the latter replaced their own hand. This did not occur when the rubber hand was rotated by 180° or replaced by a piece of wood. This illusion indicates that our brain does not build a sense of self in a merely reactive way, via perceptual correlations; rather it generates predictions on what may or may not belong to itself.     

Vibrotactile Sensory Substitution Elicits Feeling of Ownership of an Alien Hand

Tactile feedback plays a key role in the attribution of a limb to the self and in the motor control of grasping and manipulation. However, due to technological limits, current prosthetic hands do not provide amputees with cutaneous touch feedback. Recent findings showed that amputees can be tricked into experiencing an alien rubber hand as part of their own body, by applying synchronous touches to the stump which is out of view, and to the rubber hand in full view. It was suggested that similar effects could be achieved by using a prosthesis with touch sensors that provides synchronous cutaneous feedback through an array of tactile stimulators on the stump. Such a prosthesis holds the potential to be easily incorporated within one’s body scheme, because it would reproduce the perceptual illusion in everyday usage. We propose to use sensory substitution – specifically vibrotactile – to address this issue, as current haptic technology is still too bulky and inefficient. In this basic study we addressed the fundamental question of whether visuo-tactile modality mismatch promotes self-attribution of a limb, and to what extent compared to a modality-matched paradigm, on normally-limbed subjects. We manipulated visuo-tactile stimulations, comprising combinations of modality matched, modality mismatched, synchronous and asynchronous stimulations, in a set of experiments fashioned after the Rubber Hand Illusion. Modality mismatched stimulation was provided using a keypad-controlled vibrotactile display. Results from three independent measures of embodiment (questionnaires, pointing tests and skin conductance responses) indicate that vibrotactile sensory substitution can be used to induce self-attribution of a rubber hand when synchronous but modality-conflicting visuo-tactile stimulation is delivered to the biological finger pads and to the equivalent rubber hand phalanges.     

“I Feel Contaminated in My Fake Hand”: Obsessive-Compulsive-Disorder like Disgust Sensations Arise from Dummy during Rubber Hand Illusion

Despite its theoretical and clinical interest, there are no experimental studies exploring obsessive-compulsive disorder (OCD)-like disgust sensations through using somatosensory illusions. Such illusions provide important clues to the nature and limits of multisensory integration and how the brain constructs body image; and may potentially inform novel therapies. One such effect is the rubber hand illusion (RHI) in which tactile sensations are referred to a rubber hand; if the experimenter simultaneously strokes a subject’s occluded hand together with a visible fake hand, the subject starts experiencing the touch sensations as arising from the dummy. In this study, we explore whether OCD-like disgust may result from contamination of a dummy hand during the RHI; suggesting a possible integration of somatosensory and limbic inputs in the construction of body image. We predicted that participants would experience sensations of disgust, when placing a disgust stimulus (fake feces, vomit or blood) on the dummy hand after establishing the RHI. We found that 9 out of 11 participants experienced greater disgust during the synchronous condition (real hidden hand and fake hand are stroked in synchrony) compared to the asynchronous control condition (real hidden hand and fake hand are stroked in asynchrony); and on average such disgust was significantly greater during the synchronous condition compared to the asynchronous control condition, Z = 2.7, p = .008. These results argue against a strictly hierarchical modular approach to brain function and suggest that a four-way multisensory interaction occurs between vision, touch, proprioception on the one hand and primal emotions like disgust on the other. These findings may inform novel clinical approaches for OCD; that is, contaminating a dummy during the RHI could possibly be used as part of an in-vivo exposure-intervention for OCD.     

Bodily illusions modulate tactile perception

Touch differs from other exteroceptive senses in that the body itself forms part of the tactile percept. Interactions between proprioception and touch provide a powerful way to investigate the implicit body representation underlying touch. Here, we demonstrate that an intrinsic primary quality of a tactile object, for example its size, is directly affected by the perceived size of the body part touching it. We elicited proprioceptive illusions that the left index finger was either elongating or shrinking by vibrating the biceps or triceps tendon of the right arm while subjects grasped the tip of their left index finger. Subjects estimated the distance between two simultaneous tactile contacts on the left finger during tendon vibration. We found that tactile distances feel bigger when the touched body part feels elongated. Control tests showed that the modulation of touch was linked to the perceived index-finger size induced by tendon vibration. Vibrations that did not produce proprioceptive illusion had no effect on touch. Our results show that the perception of tactile objects is referenced to an implicit body representation and that proprioception contributes to this body representation. We also provide, for the first time, a quantitative, implicit measure of distortions of body size.     

Are Movements Necessary for the Sense of Body Ownership? Evidence from the Rubber Hand Illusion in Pure Hemiplegic Patients

A question still debated within cognitive neuroscience is whether signals present during actions significantly contribute to the emergence of human’s body ownership. In the present study, we aimed at answer this question by means of a neuropsychological approach. We administered the classical rubber hand illusion paradigm to a group of healthy participants and to a group of neurological patients affected by a complete left upper limb hemiplegia, but without any propriceptive/tactile deficits. The illusion strength was measured both subjectively (i.e., by a self-report questionnaire) and behaviorally (i.e., the location of one’s own hand is shifted towards the rubber hand). We aimed at examining whether, and to which extent, an enduring absence of movements related signals affects body ownership. Our results showed that patients displayed, respect to healthy participants, stronger illusory effects when the left (affected) hand was stimulated and no effects when the right (unaffected) hand was stimulated. In other words, hemiplegics had a weaker/more flexible sense of body ownership for the affected hand, but an enhanced/more rigid one for the healthy hand. Possible interpretations of such asymmetrical distribution of body ownership, as well as limits of our results, are discussed. Broadly speaking, our findings suggest that the alteration of the normal flow of signals present during movements impacts on human’s body ownership. This in turn, means that movements have a role per se in developing and maintaining a coherent body ownership.     

Explaining Away the Body: Experiences of Supernaturally Caused Touch and Touch on Non-Hand Objects within the Rubber Hand Illusion

Background

In rubber hand illusions and full body illusions, touch sensations are projected to non-body objects such as rubber hands, dolls or virtual bodies. The robustness, limits and further perceptual consequences of such illusions are not yet fully explored or understood. A number of experiments are reported that test the limits of a variant of the rubber hand illusion.

Methodology/Principal Findings

A variant of the rubber hand illusion is explored, in which the real and foreign hands are aligned in personal space. The presence of the illusion is ascertained with participants'' scores and temperature changes of the real arm. This generates a basic illusion of touch projected to a foreign arm. Participants are presented with further, unusual visuotactile stimuli subsequent to onset of the basic illusion. Such further visuotactile stimulation is found to generate very unusual experiences of supernatural touch and touch on a non-hand object. The finding of touch on a non-hand object conflicts with prior findings, and to resolve this conflict a further hypothesis is successfully tested: that without prior onset of the basic illusion this unusual experience does not occur.

Conclusions/Significance

A rubber hand illusion is found that can arise when the real and the foreign arm are aligned in personal space. This illusion persists through periods of no tactile stimulation and is strong enough to allow very unusual experiences of touch felt on a cardboard box and experiences of touch produced at a distance, as if by supernatural causation. These findings suggest that one''s visual body image is explained away during experience of the illusion and they may be of further importance to understanding the role of experience in delusion formation. The findings of touch on non-hand objects may help reconcile conflicting results in this area of research. In addition, new evidence is provided that relates to the recently discovered psychologically induced temperature changes that occur during the illusion.     

Face or hand,not both: perceptual correlates of reafferentation in a former amputee

The topography of the somatosensory maps of our body can be largely shaped by alterations of peripheral sensory inputs. Following hand amputation, the hand cortical territory becomes responsive to facial cutaneous stimulation. Amputation-induced remapping, however, reverses after transplantation, as the grafted hand (re)gains its sensorimotor representation. Here, we investigate hand tactile perception in a former amputee by touching either grafted hand singly or in combination with another body part. The results showed that tactile sensitivity recovered rapidly, being remarkably good 5 months after transplant. In the right grafted hand, however, the newly acquired somatosensory awareness was strikingly hampered when the ipsilateral face was touched simultaneously, i.e., right face perception extinguished right hand perception. Ipsilateral face-hand extinction was present in the formerly dominant right hand 5 months after transplant and eventually disappeared 6 months afterwards. Control conditions' results showed that right hand tactile awareness was not extinguished either by contralateral left face and left hand stimulation or ipsilateral stimulation of the arm, which is bodily close to, but cortically far from, the hand. We suggest that ipsilateral face-hand extinction is a perceptual counterpart of the remapping that occurs after allograft and eyewitnesses the inherently competitive nature of sensory representations.     

The illusion of owning a third arm

Could it be possible that, in the not-so-distant future, we will be able to reshape the human body so as to have extra limbs? A third arm helping us out with the weekly shopping in the local grocery store, or an extra artificial limb assisting a paralysed person? Here we report a perceptual illusion in which a rubber right hand, placed beside the real hand in full view of the participant, is perceived as a supernumerary limb belonging to the participant''s own body. This effect was supported by questionnaire data in conjunction with physiological evidence obtained from skin conductance responses when physically threatening either the rubber hand or the real one. In four well-controlled experiments, we demonstrate the minimal required conditions for the elicitation of this “supernumerary hand illusion”. In the fifth, and final experiment, we show that the illusion reported here is qualitatively different from the traditional rubber hand illusion as it is characterised by less disownership of the real hand and a stronger feeling of having two right hands. These results suggest that the artificial hand ‘borrows’ some of the multisensory processes that represent the real hand, leading to duplication of touch and ownership of two right arms. This work represents a major advance because it challenges the traditional view of the gross morphology of the human body as a fundamental constraint on what we can come to experience as our physical self, by showing that the body representation can easily be updated to incorporate an additional limb.     

Dissociation between Emotional Remapping of Fear and Disgust in Alexithymia

There is growing evidence that individuals are able to understand others’ emotions because they “embody” them, i.e., re-experience them by activating a representation of the observed emotion within their own body. One way to study emotion embodiment is provided by a multisensory stimulation paradigm called emotional visual remapping of touch (eVRT), in which the degree of embodiment/remapping of emotions is measured as enhanced detection of near-threshold tactile stimuli on one’s own face while viewing different emotional facial expressions. Here, we measured remapping of fear and disgust in participants with low (LA) and high (HA) levels of alexithymia, a personality trait characterized by a difficulty in recognizing emotions. The results showed that fear is remapped in LA but not in HA participants, while disgust is remapped in HA but not in LA participants. To investigate the hypothesis that HA might exhibit increased responses to emotional stimuli producing a heightened physical and visceral sensations, i.e., disgust, in a second experiment we investigated participants’ interoceptive abilities and the link between interoception and emotional modulations of VRT. The results showed that participants’ disgust modulations of VRT correlated with their ability to perceive bodily signals. We suggest that the emotional profile of HA individuals on the eVRT task could be related to their abnormal tendency to be focalized on their internal bodily signals, and to experience emotions in a “physical” way. Finally, we speculated that these results in HA could be due to a enhancement of insular activity during the perception of disgusted faces.     

Tactile Gap Detection Deteriorates during Bimanual Symmetrical Movements under Mirror Visual Feedback

It has been suggested that incongruence between signals for motor intention and sensory input can cause pain and other sensory abnormalities. This claim is supported by reports that moving in an environment of induced sensorimotor conflict leads to elevated pain and sensory symptoms in those with certain painful conditions. Similar procedures can lead to reports of anomalous sensations in healthy volunteers too. In the present study, we used mirror visual feedback to investigate the effects of sensorimotor incongruence on responses to stimuli that arise from sources external to the body, in particular, touch. Incongruence between the sensory and motor signals for the right arm was manipulated by having the participants make symmetrical or asymmetrical movements while watching a reflection of their left arm in a parasagittal mirror, or the left hand surface of a similarly positioned opaque board. In contrast to our prediction, sensitivity to the presence of gaps in tactile stimulation of the right forearm was not reduced when participants made asymmetrical movements during mirror visual feedback, as compared to when they made symmetrical or asymmetrical movements with no visual feedback. Instead, sensitivity was reduced when participants made symmetrical movements during mirror visual feedback relative to the other three conditions. We suggest that small discrepancies between sensory and motor information, as they occur during mirror visual feedback with symmetrical movements, can impair tactile processing. In contrast, asymmetrical movements with mirror visual feedback may not impact tactile processing because the larger discrepancies between sensory and motor information may prevent the integration of these sources of information. These results contrast with previous reports of anomalous sensations during exposure to both low and high sensorimotor conflict, but are nevertheless in agreement with a forward model interpretation of perceptual modulations during goal directed movement.     

The Importance of Synchrony and Temporal Order of Visual and Tactile Input for Illusory Limb Ownership Experiences – An fMRI Study Applying Virtual Reality

In the so-called rubber hand illusion, synchronous visuotactile stimulation of a visible rubber hand together with one''s own hidden hand elicits ownership experiences for the artificial limb. Recently, advanced virtual reality setups were developed to induce a virtual hand illusion (VHI). Here, we present functional imaging data from a sample of 25 healthy participants using a new device to induce the VHI in the environment of a magnetic resonance imaging (MRI) system. In order to evaluate the neuronal robustness of the illusion, we varied the degree of synchrony between visual and tactile events in five steps: in two conditions, the tactile stimulation was applied prior to visual stimulation (asynchrony of −300 ms or −600 ms), whereas in another two conditions, the tactile stimulation was applied after visual stimulation (asynchrony of +300 ms or +600 ms). In the fifth condition, tactile and visual stimulation was applied synchronously. On a subjective level, the VHI was successfully induced by synchronous visuotactile stimulation. Asynchronies between visual and tactile input of ±300 ms did not significantly diminish the vividness of illusion, whereas asynchronies of ±600 ms did. The temporal order of visual and tactile stimulation had no effect on VHI vividness. Conjunction analyses of functional MRI data across all conditions revealed significant activation in bilateral ventral premotor cortex (PMv). Further characteristic activation patterns included bilateral activity in the motion-sensitive medial superior temporal area as well as in the bilateral Rolandic operculum, suggesting their involvement in the processing of bodily awareness through the integration of visual and tactile events. A comparison of the VHI-inducing conditions with asynchronous control conditions of ±600 ms yielded significant PMv activity only contralateral to the stimulation site. These results underline the temporal limits of the induction of limb ownership related to multisensory body-related input.     

Negative Body Image Associated with Changes in the Visual Body Appearance Increases Pain Perception

Changing the visual body appearance by use of as virtual reality system, funny mirror, or binocular glasses has been reported to be helpful in rehabilitation of pain. However, there are interindividual differences in the analgesic effect of changing the visual body image. We hypothesized that a negative body image associated with changing the visual body appearance causes interindividual differences in the analgesic effect although the relationship between the visual body appearance and analgesic effect has not been clarified. We investigated whether a negative body image associated with changes in the visual body appearance increased pain. Twenty-five healthy individuals participated in this study. To evoke a negative body image, we applied the method of rubber hand illusion. We created an “injured rubber hand” to evoke unpleasantness associated with pain, a “hairy rubber hand” to evoke unpleasantness associated with embarrassment, and a “twisted rubber hand” to evoke unpleasantness associated with deviation from the concept of normality. We also created a “normal rubber hand” as a control. The pain threshold was measured while the participant observed the rubber hand using a device that measured pain caused by thermal stimuli. Body ownership experiences were elicited by observation of the injured rubber hand and hairy rubber hand as well as the normal rubber hand. Participants felt more unpleasantness by observing the injured rubber hand and hairy rubber hand than the normal rubber hand and twisted rubber hand (p<0.001). The pain threshold was lower under the injured rubber hand condition than with the other conditions (p<0.001). We conclude that a negative body appearance associated with pain can increase pain sensitivity.     

The Right Planum Temporale Is Involved in Stimulus-Driven,Auditory Attention – Evidence from Transcranial Magnetic Stimulation

It is well known that the planum temporale (PT) area in the posterior temporal lobe carries out spectro-temporal analysis of auditory stimuli, which is crucial for speech, for example. There are suggestions that the PT is also involved in auditory attention, specifically in the discrimination and selection of stimuli from the left and right ear. However, direct evidence is missing so far. To examine the role of the PT in auditory attention we asked fourteen participants to complete the Bergen Dichotic Listening Test. In this test two different consonant-vowel syllables (e.g., “ba” and “da”) are presented simultaneously, one to each ear, and participants are asked to verbally report the syllable they heard best or most clearly. Thus attentional selection of a syllable is stimulus-driven. Each participant completed the test three times: after their left and right PT (located with anatomical brain scans) had been stimulated with repetitive transcranial magnetic stimulation (rTMS), which transiently interferes with normal brain functioning in the stimulated sites, and after sham stimulation, where participants were led to believe they had been stimulated but no rTMS was applied (control). After sham stimulation the typical right ear advantage emerged, that is, participants reported relatively more right than left ear syllables, reflecting a left-hemispheric dominance for language. rTMS over the right but not left PT significantly reduced the right ear advantage. This was the result of participants reporting more left and fewer right ear syllables after right PT stimulation, suggesting there was a leftward shift in stimulus selection. Taken together, our findings point to a new function of the PT in addition to auditory perception: particularly the right PT is involved in stimulus selection and (stimulus-driven), auditory attention.     

Seeing the body produces limb-specific modulation of skin temperature

Vision of the body, even when non-informative about stimulation, affects somatosensory processing. We investigated whether seeing the body also modulates autonomic control in the periphery by measuring skin temperature while manipulating vision. Using a mirror box, the skin temperature was measured from left hand dorsum while participants: (i) had the illusion of seeing their left hand, (ii) had the illusion of seeing an object at the same location or (iii) looked directly at their contralateral right hand. Skin temperature of the left hand increased when participants had the illusion of directly seeing that hand but not in the other two view conditions. In experiment 2, participants viewed directly their left or right hand, or the box while we recorded both hand dorsum temperatures. Temperature increased in the viewed hand but not the contralateral hand. These results show that seeing the body produces limb-specific modulation of thermal regulation.     

Response of thoracic interneurons to tactile stimulation in the cockroach,Periplaneta americana

Receåfindings indicate that cockroaches escape in response to tactile stimulation as well as they do in response to the classic wind puff stimulus. The thoracic interneurons that receive inputs from ventral giant interneurons also respond to tactile stimulation and therefore, represent a potential site of convergence between wind and tactile stimulation as well as other sensory modalities. In this article, we characterize the tactile response of these interneurons, which are referred to as type-A thoracic interneurons (TI_As). In response to tactile stimulation of the body cuticle, TI_As typically respond with a short latency biphasic depolarization which often passes threshold for action potentials. The biphasic response is not typical of responses to wind stimulation nor of tactile stimulation of the antennae. It is also not seen in tactile responses of thoracic interneurons that are not part of the TI_A group. The responses of individual TI_As to stimulation of various body locations were mapped. The left-right directional properties of TI_As are consistent with their responses to wind puffs from various different directions. Cells that respond equally well to wind from the left and right side also respond equally well to tactile stimuli on the left and right side of the animal's body. In contrast, cells that are biased to wind on one side are also biased to tactile stimulation on the same side. In general, tactile responses directed at body cuticle are phasic rather than tonic, occurring both when the tactile stimulator is depressed and released. The response reflects stimulus strength and follows repeated stimulation quite well. However, the first phase of the biphasic response is more robust during high-frequency stimulation than the second phase. TI_As also respond to antennal stimulation. However, here the response characteristics are complicated by the fact that movement of either antenna evokes descending activity in both left and right thoracic connectives. The data suggest that the TI_As make up a multimodal site of sensory convergence that is capable of generating an oriental escape turn in response to any one of several sensory cues. 1994 John Wiley & Sons, Inc.     

The Crossmodal Congruency Task as a Means to Obtain an Objective Behavioral Measure in the Rubber Hand Illusion Paradigm

The rubber hand illusion (RHI) is a popular experimental paradigm. Participants view touch on an artificial rubber hand while the participants'' own hidden hand is touched. If the viewed and felt touches are given at the same time then this is sufficient to induce the compelling experience that the rubber hand is one''s own hand. The RHI can be used to investigate exactly how the brain constructs distinct body representations for one''s own body. Such representations are crucial for successful interactions with the external world. To obtain a subjective measure of the RHI, researchers typically ask participants to rate statements such as "I felt as if the rubber hand were my hand". Here we demonstrate how the crossmodal congruency task can be used to obtain an objective behavioral measure within this paradigm.The variant of the crossmodal congruency task we employ involves the presentation of tactile targets and visual distractors. Targets and distractors are spatially congruent (i.e. same finger) on some trials and incongruent (i.e. different finger) on others. The difference in performance between incongruent and congruent trials - the crossmodal congruency effect (CCE) - indexes multisensory interactions. Importantly, the CCE is modulated both by viewing a hand as well as the synchrony of viewed and felt touch which are both crucial factors for the RHI.The use of the crossmodal congruency task within the RHI paradigm has several advantages. It is a simple behavioral measure which can be repeated many times and which can be obtained during the illusion while participants view the artificial hand. Furthermore, this measure is not susceptible to observer and experimenter biases. The combination of the RHI paradigm with the crossmodal congruency task allows in particular for the investigation of multisensory processes which are critical for modulations of body representations as in the RHI.     

The Human Touch: Skin Temperature during the Rubber Hand Illusion in Manual and Automated Stroking Procedures

A difference in skin temperature between the hands has been identified as a physiological correlate of the rubber hand illusion (RHI). The RHI is an illusion of body ownership, where participants perceive body ownership over a rubber hand if they see it being stroked in synchrony with their own occluded hand. The current study set out to replicate this result, i.e., psychologically induced cooling of the stimulated hand using an automated stroking paradigm, where stimulation was delivered by a robot arm (PHANToM^TM force-feedback device). After we found no evidence for hand cooling in two experiments using this automated procedure, we reverted to a manual stroking paradigm, which is closer to the one employed in the study that first produced this effect. With this procedure, we observed a relative cooling of the stimulated hand in both the experimental and the control condition. The subjective experience of ownership, as rated by the participants, by contrast, was strictly linked to synchronous stroking in all three experiments. This implies that hand-cooling is not a strict correlate of the subjective feeling of hand ownership in the RHI. Factors associated with the differences between the two designs (differences in pressure of tactile stimulation, presence of another person) that were thus far considered irrelevant to the RHI appear to play a role in bringing about this temperature effect.     

Decreased Peripheral and Central Responses to Acupuncture Stimulation following Modification of Body Ownership

Acupuncture stimulation increases local blood flow around the site of stimulation and induces signal changes in brain regions related to the body matrix. The rubber hand illusion (RHI) is an experimental paradigm that manipulates important aspects of bodily self-awareness. The present study aimed to investigate how modifications of body ownership using the RHI affect local blood flow and cerebral responses during acupuncture needle stimulation. During the RHI, acupuncture needle stimulation was applied to the real left hand while measuring blood microcirculation with a LASER Doppler imager (Experiment 1, N = 28) and concurrent brain signal changes using functional magnetic resonance imaging (fMRI; Experiment 2, N = 17). When the body ownership of participants was altered by the RHI, acupuncture stimulation resulted in a significantly lower increase in local blood flow (Experiment 1), and significantly less brain activation was detected in the right insula (Experiment 2). This study found changes in both local blood flow and brain responses during acupuncture needle stimulation following modification of body ownership. These findings suggest that physiological responses during acupuncture stimulation can be influenced by the modification of body ownership.