Arvid Guterstam

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 participants 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.

The invisible hand illusion: Multisensory integration leads to the embodiment of a discrete volume of empty space

The dynamic integration of signals from different sensory modalities plays a key role in bodily self-perception. When visual information is used in the multisensory process of localizing and identifying ones own limbs, the sight of a body part often plays a dominant role. For example, it has repeatedly been shown that a viewed object must resemble a humanoid body part to permit illusory self-attribution of that object. Here, we report a perceptual illusion that challenges these assumptions by demonstrating that healthy (nonamputated) individuals can refer somatic sensations to a discrete volume of empty space and experience having an invisible hand. In 10 behavioral and one fMRI experiment, we characterized the perceptual rules and multisensory brain mechanisms that produced this “invisible hand illusion.” Our behavioral results showed that the illusion depends on visuotactile-proprioceptive integration that obeys key spatial and temporal multisensory rules confined to near-personal space. The fMRI results associate the illusion experience with increased activity in regions related to the integration of multisensory body-related signals, most notably the bilateral ventral premotor, intraparietal, and cerebellar cortices. We further showed that a stronger feeling of having an invisible hand is associated with a higher degree of effective connectivity between the intraparietal and ventral premotor cortices. These findings demonstrate that the integration of temporally and spatially congruent multisensory signals in a premotor-intraparietal circuit is sufficient to redefine the spatial boundaries of the bodily self, even when visual information directly contradicts the presence of a physical limb at the location of the perceived illusory hand.

Disintegration of Multisensory Signals from the Real Hand Reduces Default Limb Self-Attribution: An fMRI Study

The perception of our limbs in space is built upon the integration of visual, tactile, and proprioceptive signals. Accumulating evidence suggests that these signals are combined in areas of premotor, parietal, and cerebellar cortices. However, it remains to be determined whether neuronal populations in these areas integrate hand signals according to basic temporal and spatial congruence principles of multisensory integration. Here, we developed a setup based on advanced 3D video technology that allowed us to manipulate the spatiotemporal relationships of visuotactile (VT) stimuli delivered on a healthy human participants real hand during fMRI and investigate the ensuing neural and perceptual correlates. Our experiments revealed two novel findings. First, we found responses in premotor, parietal, and cerebellar regions that were dependent upon the spatial and temporal congruence of VT stimuli. This multisensory integration effect required a simultaneous match between the seen and felt postures of the hand, which suggests that congruent visuoproprioceptive signals from the upper limb are essential for successful VT integration. Second, we observed that multisensory conflicts significantly disrupted the default feeling of ownership of the seen real limb, as indexed by complementary subjective, psychophysiological, and BOLD measures. The degree to which self-attribution was impaired could be predicted from the attenuation of neural responses in key multisensory areas. These results elucidate the neural bases of the integration of multisensory hand signals according to basic spatiotemporal principles and demonstrate that the disintegration of these signals leads to “disownership” of the seen real hand.

Illusory ownership of an invisible body reduces autonomic and subjective social anxiety responses

What is it like to be invisible? This question has long fascinated man and has been the central theme of many classic literary works. Recent advances in materials science suggest that invisibility cloaking of the human body may be possible in the not-so-distant future. However, it remains unknown how invisibility affects body perception and embodied cognition. To address these questions, we developed a perceptual illusion of having an entire invisible body. Through a series of experiments, we characterized the multisensory rules that govern the elicitation of the illusion and show that the experience of having an invisible body reduces the social anxiety response to standing in front of an audience. This study provides an experimental model of what it is like to be invisible and shows that this experience affects bodily self-perception and social cognition.

Ownership of an artificial limb induced by electrical brain stimulation

Significance Creating a prosthetic device that feels like one’s own limb is a major challenge in applied neuroscience. We show that ownership of an artificial hand can be induced via electrical stimulation of the hand somatosensory cortex in synchrony with touches applied to a prosthetic hand in full view. These findings suggest that the human brain can integrate “natural” visual input and direct cortical-somatosensory stimulation to create the multisensory perception that an artificial limb belongs to one’s own body. Replacing the function of a missing or paralyzed limb with a prosthetic device that acts and feels like one’s own limb is a major goal in applied neuroscience. Recent studies in nonhuman primates have shown that motor control and sensory feedback can be achieved by connecting sensors in a robotic arm to electrodes implanted in the brain. However, it remains unknown whether electrical brain stimulation can be used to create a sense of ownership of an artificial limb. In this study on two human subjects, we show that ownership of an artificial hand can be induced via the electrical stimulation of the hand section of the somatosensory (SI) cortex in synchrony with touches applied to a rubber hand. Importantly, the illusion was not elicited when the electrical stimulation was delivered asynchronously or to a portion of the SI cortex representing a body part other than the hand, suggesting that multisensory integration according to basic spatial and temporal congruence rules is the underlying mechanism of the illusion. These findings show that the brain is capable of integrating “natural” visual input and direct cortical-somatosensory stimulation to create the multisensory perception that an artificial limb belongs to one’s own body. Thus, they serve as a proof of concept that electrical brain stimulation can be used to “bypass” the peripheral nervous system to induce multisensory illusions and ownership of artificial body parts, which has important implications for patients who lack peripheral sensory input due to spinal cord or nerve lesions.

Decoding illusory self-location from activity in the human hippocampus

Decades of research have demonstrated a role for the hippocampus in spatial navigation and episodic and spatial memory. However, empirical evidence linking hippocampal activity to the perceptual experience of being physically located at a particular place in the environment is lacking. In this study, we used a multisensory out-of-body illusion to perceptually ‘teleport’ six healthy participants between two different locations in the scanner room during high-resolution functional magnetic resonance imaging (fMRI). The participants were fitted with MRI-compatible head-mounted displays that changed their first-person visual perspective to that of a pair of cameras placed in one of two corners of the scanner room. To elicit the illusion of being physically located in this position, we delivered synchronous visuo-tactile stimulation in the form of an object moving toward the cameras coupled with touches applied to the participant’s chest. Asynchronous visuo-tactile stimulation did not induce the illusion and served as a control condition. We found that illusory self-location could be successfully decoded from patterns of activity in the hippocampus in all of the participants in the synchronous (P < 0.05) but not in the asynchronous condition (P > 0.05). At the group-level, the decoding accuracy was significantly higher in the synchronous than in the asynchronous condition (P = 0.012). These findings associate hippocampal activity with the perceived location of the bodily self in space, which suggests that the human hippocampus is involved not only in spatial navigation and memory but also in the construction of our sense of bodily self-location.

Tool use changes the spatial extension of the magnetic touch illusion

Characterizing the brain mechanisms that allow humans to use tools to interact with the environment is a major goal in neuroscience. It has been proposed that handheld tools are incorporated into the multisensory representation of the body and its surrounding (peripersonal) space, underlying our remarkable tool use ability. One single-cell recording study in tool-using monkeys provided qualitative support for this hypothesis, and the results from a vast number of human studies employing different experimental paradigms have been ambiguous. Here, we made use of the recently reported magnetic touch illusion—a perceptual correlate of peripersonal space—to examine the effect of tool use on the representation of visuotactile peripersonal space. The results showed that active tool use leads to an extension of the “illusion volume” around the entire length of a tool, which was significantly greater compared with a manual control task. These findings support the notion that the multisensory representation of peripersonal space is extended to incorporate handheld tools and provide a three-dimensional estimation of this remapping process.