当代意识科学面临着双重挑战：既要说明意识的机制，同时要说明与之相关的哲学问题。达马西奥从生命自我保存和自我生产的生物价值出发，将意识视为生命有机体在演化进程中形成的一种高级、奇妙的调节和管理生命的方式。他从这个角度提出了一种意识机制的演化学说——具有自我感的核心自我是通过表征被客体改变的原自我的状态而建构起来的，它是一种二阶映射。他的意识理论不可避免地要涉及对诸如心一身问题、自我问题、认识论问题等传统哲学主题的见解和主张。

身体意象是个体在意识层面对与身体有关的知觉、态度及信念,它的稳定性与可塑性一直存在争论.本研究使用方位追踪仪、数据手套及投影仪,通过2个实验考查虚拟手错觉中距离参照系对身体拥有感的影响.实验1在虚拟环境中采用经典橡胶手错觉的变式,考察同步性和虚拟手位置对拥有感的影响;实验2引入距离参照系,考察在不同参照条件下被试对同一个位置的虚拟手在不同同步情况下的拥有感错觉.研究结果发现:(1)无论是同步性还是距离都会对虚拟手错觉中的拥有感产生影响,即同步比不同步条件下、距离近比距离远条件下拥有感显著;(2)不同的距离参照系对同一位置的拥有感体验影响差异显著,先近后远的呈现方式会减弱错觉而先远后近的呈现方式则会加强错觉.这一发现与身体拥有感是受稳定的身体意象调节的假设并不一致.

Recognising and representing one's self as distinct from others is a fundamental component of self-awareness. However, current theories of self-recognition are not embedded within global theories of cortical function and therefore fail to provide a compelling explanation of how the self is processed. We present a theoretical account of the neural and computational basis of self-recognition that is embedded within the free-energy account of cortical function. In this account one's body is processed in a Bayesian manner as the most likely to be “me”. Such probabilistic representation arises through the integration of information from hierarchically organised unimodal systems in higher-level multimodal areas. This information takes the form of bottom-up “surprise” signals from unimodal sensory systems that are explained away by top-down processes that minimise the level of surprise across the brain. We present evidence that this theoretical perspective may account for the findings of psychological and neuroimaging investigations into self-recognition and particularly evidence that representations of the self are malleable, rather than fixed as previous accounts of self-recognition might suggest.

Abstract Subjects perceived touch sensations as arising from a table (or a rubber hand) when both the table (or the rubber hand) and their own real hand were repeatedly tapped and stroked in synchrony with the real hand hidden from view. If the table or rubber hand was then 'injured', subjects displayed a strong skin conductance response (SCR) even though nothing was done to the real hand. Sensations could even be projected to anatomically impossible locations. The illusion was much less vivid, as indicated by subjective reports and SCR, if the real hand was simultaneously visible during stroking, or if the real hand was hidden but touched asynchronously. The fact that the illusion could be significantly diminished when the real hand was simultaneously visible suggests that the illusion and associated SCRs were due to perceptual assimilation of the table (or rubber hand) into one's body image rather than associative conditioning. These experiments demonstrate the malleability of body image and the brain's remarkable capacity for detecting statistical correlations in the sensory input.

Cerebral damage may induce a delusional belief so that patients claim that their limbs contralateral to the side of the lesion belong to someone else (somatoparaphrenia). This disorder, which is not due to a general , is frequently accompanied by the inability to feel tactile sensations in the 'non-belonging' part of the body. We report the unique case of a patient with somatoparaphrenia in whom dense tactile imperception in the left hand dramatically recovered when she was instructed to report touches delivered to her niece's hand, rather than to her own hand. We suggest that, through this verbal instruction, the mismatch between the patient's belief about the ownership of her left hand and her ability to perceive touch on it was transiently recomposed. This is evidence that apparently elementary deficits, such as hemianesthesia, and selective delusional , such as somatoparaphrenia, may both originate from an impairment of the body image.

Botvinick M, Cohen J.

Behavioral studies in humans and electrophysiological recordings in nonhuman primates have suggested the existence of a specific representation of the space immediately surrounding the body. In macaques, neurons that have visual receptive fields limited to a region of space close around a body part have been found in premotor and parietal areas. These cells are hypothesized to encode the location of external objects in coordinate systems that are centered on individual body parts. In the present study, we used an fMRI adaptation paradigm on healthy participants to reveal areas in the anterior part of the intraparietal sulcus, the inferior parietal lobe (supramarginal gyrus), and the dorsal and ventral portions of the premotor cortex that exhibit selective BOLD adaptation to an object moving near the right hand. Crucially, these areas did not manifest adaptation if the stimulus was presented in far space (100 cm) or when the hand was retracted from the object. This hand-centered selectivity could not be detected when a traditional fMRI analysis approach was used. These findings are important as they provide the most conclusive neuroimaging evidence to date for a representation of near-personal space in the human brain. They also demonstrate a selective mechanism implemented by human perihand neurons in the premotor and posterior parietal areas and add to earlier findings from humans and nonhuman primates.

Cognitive neuroscience investigations of self-experience have mainly focused on the mental attribution of features to the self (self-related processing). In this paper, we highlight another fundamental, yet neglected, aspect of self-experience, that of being an agent. We propose that this aspect of self-experience depends on self-specifying processes, ones that implicitly specify the self by implementing a functional self/non-self distinction in perception, action, cognition and emotion. We describe two paradigmatic cases – sensorimotor integration and homeostatic regulation – and use the principles from these cases to show how cognitive control, including emotion regulation, is also self-specifying. We argue that externally directed, attention-demanding tasks, rather than suppressing self-experience, give rise to the self-experience of being a cognitive–affective agent. We conclude with directions for experimental work based on our framework.

The sense that I am the author of my own actions, including the ability to distinguish my own from other people鈥檚 actions, is a fundamental building block of our sense of self, on the one hand, and successful social interactions, on the other. Using cognitive neuroscience techniques, researchers have attempted to elucidate the functional basis of this intriguing phenomenon, also trying to explain pathological abnormalities of action awareness in certain psychiatric and neurological disturbances. Recent conceptual, technological and methodological advances suggest several interesting and necessary new leads for future research on the neuroscience of agency. Here I will describe new frontiers for the field such as the need for novel and multifactorial paradigms, anatomically plausible network models for the sense of agency, investigations of the temporal dynamics during agentic processing and ecologically valid virtual reality applications.

Abstract The sense of agency is a central aspect of human self-consciousness and refers to the experience of oneself as the agent of one's own actions. Several different cognitive theories on the sense of agency have been proposed implying divergent empirical approaches and results, especially with respect to neural correlates. A multifactorial and multilevel model of the sense of agency may provide the most constructive framework for integrating divergent theories and findings, meeting the complex nature of this intriguing phenomenon.

Alien hand syndrome (AHS) is an involuntary and rare neurological disorder emerges at upper extremity. AHS is a disconnection syndrome with the symptoms of losing sense of agency and sense of ownershi

The feeling of body ownership is a fundamental aspect of self-consciousness. The underlying neural mechanisms can be studied by using the illusion where a person is made to feel that a rubber hand is his or her own hand by brushing the person's hidden real hand and synchronously brushing the artificial hand that is in full view. Here we show that threat to the rubber hand can induce a similar level of activity in the brain areas associated with anxiety and interoceptive awareness (insula and anterior cingulate cortex) as when the person's real hand is threatened. We further show that the stronger the feeling of ownership of the artificial hand, the stronger the threat-evoked neuronal responses in the areas reflecting anxiety. Furthermore, across subjects, activity in multisensory areas reflecting ownership predicted the activity in the interoceptive system when the hand was under threat. Finally, we show that there is activity in medial wall motor areas, reflecting an urge to withdraw the artificial hand when it is under threat. These findings suggest that artificial limbs can evoke the same feelings as real limbs and provide objective neurophysiological evidence that the rubber hand is fully incorporated into the body. These findings are of fundamental importance because they suggest that the feeling of body ownership is associated with changes in the interoceptive systems.

Abstract It has been hypothesized that an internal model is involved in controlling and recognizing one's own actions (action attribution). This results from a comparison process between the predicted sensory feedback of the action and its real sensory consequences. The aim of the present study is to distinguish the respective importance of two action parameters (time and direction) on such an attribution judgment. We used a device that allows introduction of discordance between the movements actually performed and the sensory feedback displayed on a computer screen. Participants were asked to judge whether they were viewing 1) their own movements, 2) their own movements modified (spatially or temporally displaced), or 3) those of another agent (i.e, the experimenter). In fact, in all conditions they were only shown their own movements either unaltered or modified by varying amounts in space or time. Movements were only attributed to another agent when there was a high spatial discordance between participants' hand movements and sensory feedback. This study is the first to show that the direction of movements is a cardinal feature in action attribution, whereas temporal properties of movements play a less important role.

This article concerns the nature of evoked brain responses and the principles underlying their generation. We start with the premise that the sensory brain has evolved to represent or infer the causes of changes in its sensory inputs. The problem of inference is well formulated in statistical terms. The statistical fundaments of inference may therefore afford important constraints on neuronal implementation. By formulating the original ideas of Helmholtz on perception, in terms of modern-day statistical theories, one arrives at a model of perceptual inference and learning that can explain a remarkable range of neurobiological facts.It turns out that the problems of inferring the causes of sensory input (perceptual inference) and learning the relationship between input and cause (perceptual learning) can be resolved using exactly the same principle. Specifically, both inference and learning rest on minimizing the brain's free energy, as defined in statistical physics. Furthermore, inference and learning can proceed in a biologically plausible fashion. Cortical responses can be seen as the brain's attempt to minimize the free energy induced by a stimulus and thereby encode the most likely cause of that stimulus. Similarly, learning emerges from changes in synaptic efficacy that minimize the free energy, averaged over all stimuli encountered. The underlying scheme rests on empirical Bayes and hierarchical models of how sensory input is caused. The use of hierarchical models enables the brain to construct prior expectations in a dynamic and context-sensitive fashion. This scheme provides a principled way to understand many aspects of cortical organization and responses. The aim of this article is to encompass many apparently unrelated anatomical, physiological and psychophysical attributes of the brain within a single theoretical perspective. In terms of cortical architectures, the theoretical treatment predicts that sensory cortex should be arranged hierarchically, that connections should be reciprocal and that forward and backward connections should show a functional asymmetry (forward connections are driving, whereas backward connections are both driving and modulatory). In terms of synaptic physiology, it predicts associative plasticity and, for dynamic models, spike-timing-dependent plasticity. In terms of electrophysiology, it accounts for classical and extra classical receptive field effects and long-latency or endogenous components of evoked cortical responses. It predicts the attenuation of responses encoding prediction error with perceptual learning and explains many phenomena such as repetition suppression, mismatch negativity (MMN) and the P300 in electroencephalography. In psychophysical terms, it accounts for the behavioural correlates of these physiological phenomena, for example, priming and global precedence. The final focus of this article is on perceptual learning as measured with the MMN and the implications for empirical studies of coupling among cortical areas using evoked sensory responses.

A free-energy principle has been proposed recently that accounts for action, perception and learning. This Review looks at some key brain theories in the biological (for example, neural Darwinism) and physical (for example, information theory and optimal control theory) sciences from the free-energy perspective. Crucially, one key theme runs through each of these theories - optimization. Furthermore, if we look closely at what is optimized, the same quantity keeps emerging, namely value (expected reward, expected utility) or its complement, surprise (prediction error, expected cost). This is the quantity that is optimized under the free-energy principle, which suggests that several global brain theories might be unified within a free-energy framework.

The aim of our paper is to show that there is a sense of body that is enactive in nature and that enables to capture the most primitive sense of self. We will argue that the body is primarily given to us as source or power for action, i.e., as the variety of motor potentialities that define the horizon of the world in which we live, by populating it with things at hand to which we can be directed and with other bodies we can interact with. We will show that this sense of body as bodily self is, on the one hand, antecedent the distinction between sense of agency and sense of ownership, and, on the other, it enables and refines such distinction, providing a conceptual framework for the coherent interpretation of a variety of behavioral and neuropsychological data. We will conclude by positing that the basic experiences we entertain of our selves as bodily selves are from the very beginning driven by our interactions with other bodies as they are underpinned by the mirror mechanism.

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 participant's 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.

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.

The capacity for voluntary action is seen as essential to human nature. Yet neuroscience and behaviourist psychology have traditionally dismissed the topic as unscientific, perhaps because the mechanisms that cause actions have long been unclear. However, new research has identified networks of brain areas, including the pre-supplementary motor area, the anterior prefrontal cortex and the parietal cortex, that underlie voluntary action. These areas generate information for forthcoming actions, and also cause the distinctive conscious experience of intending to act and then controlling one's own actions. Volition consists of a series of decisions regarding whether to act, what action to perform and when to perform it. Neuroscientific accounts of voluntary action may inform debates about the nature of individual responsibility.

In adult life, people normally know what they are doing. This experience of controlling one's own actions and, through them, the course of events in the outside world is called 'sense of agency'. It forms a central feature of human experience; however, the brain mechanisms that produce the sense of agency have only recently begun to be investigated systematically. This recent progress has been driven by the development of better measures of the experience of agency, improved design of cognitive and behavioural experiments, and a growing understanding of the brain circuits that generate this distinctive but elusive experience. The sense of agency is a mental and neural state of cardinal importance in human civilization, because it is frequently altered in psychopathology and because it underpins the concept of responsibility in human societies.

Alien hand syndrome is a term used to describe a variety of rare conditions in which uncontrolled behavior or feelings of strangeness are felt in one extremity, most commonly the left hand. Etiology usually involves infarct of the right anterior or posterior cerebral arteries or cortical-basal degeneration. The medical and neuropsychological data of an elderly female who suffered a left middle cerebral artery stroke with resulting right-sided alien hand sign is presented. Neuropsychological assessment revealed declines in visual- and perceptual-based abilities and right-sided motor and sensory abilities consistent with the affected areas indicated on neuroimaging. This case demonstrates the utility of neuropsychological assessment in patients with unusual sensory/motor presentations.

Abstract Self-consciousness has mostly been approached by philosophical enquiry and not by empirical neuroscientific study, leading to an overabundance of diverging theories and an absence of data-driven theories. Using robotic technology, we achieved specific bodily conflicts and induced predictable changes in a fundamental aspect of self-consciousness by altering where healthy subjects experienced themselves to be (self-location). Functional magnetic resonance imaging revealed that temporo-parietal junction (TPJ) activity reflected experimental changes in self-location that also depended on the first-person perspective due to visuo-tactile and visuo-vestibular conflicts. Moreover, in a large lesion analysis study of neurological patients with a well-defined state of abnormal self-location, brain damage was also localized at TPJ, providing causal evidence that TPJ encodes self-location. Our findings reveal that multisensory integration at the TPJ reflects one of the most fundamental subjective feelings of humans: the feeling of being an entity localized at a position in space and perceiving the world from this position and perspective. Copyright 2011 Elsevier Inc. All rights reserved.

Abstract Recognizing oneself as the owner of a body and the agent of actions requires specific mechanisms which have been elucidated only recently. One of these mechanisms is the monitoring of signals arising from bodily movements, i.e. the central signals which contribute to the generation of the movements and the sensory signals which arise from their execution. The congruence between these two sets of signals is a strong index for determining the experiences of ownership and agency, which are the main constituents of the experience of being an independent self. This mechanism, however, does not account from the frequent cases where an intention is generated but the corresponding action is not executed. In this paper, it is postulated that such covert actions are internally simulated by activating specific cortical networks or representations of the intended actions. This process of action simulation is also extended to the observation and the recognition of actions performed or intended by other agents. The problem of disentangling representations that pertain to self-intended actions from those that pertain to actions executed or intended by others, is a critical one for attributing actions to their respective agents. Failure to recognize one's own actions and misattribution of actions may result from pathological conditions which alter the readability of these representations.

The rubber hand illusion is a perceptual illusion in which a model hand is experienced as part of one own body. In the present study we directly compared the classical illusion, based on visuotactile stimulation, with a rubber hand illusion based on active and passive movements. We examined the question of which combinations of sensory and motor cues are the most potent in inducing the illusion by subjective ratings and an objective measure (proprioceptive drift). In particular, we were interested in whether the combination of afferent and efferent signals in active movements results in the same illusion as in the purely passive modes. Our results show that the illusion is equally strong in all three cases. This demonstrates that different combinations of sensory input can lead to a very similar phenomenological experience and indicates that the illusion can be induced by any combination of multisensory information.

The ability to control external events through our own actions is a fundamental aspect of human experience. Both the subjective experience of agency, and its neural correlates, remain poorly understood. Previous studies show that the angular gyrus is activated when participants explicitly judge that they lack agency. In contrast, the positive sense of agency over external events is associated with distortions of time perception. Here, we show that the perceived interval between actions and a subsequent tone is shorter than the perceived interval between a physically comparable passive movement and a tone, replicating the ntentional binding effect reported previously. We considered this as a potential implicit marker of agency, and investigated its neural basis, by using parametric analyses to identify brain areas whose activation correlated more strongly with the perceived action-tone interval in the action condition, than in the passive condition. Small volume corrections were used to test specific hypotheses about the contribution of the angular gyrus, and of the supplementary motor area (SMA), based on previous literature. We found no correlation between angular gyrus and our temporal measure of sense of agency. In contrast, we found that a lateral, caudal region within the SMA proper was more strongly associated with the perceived action-tone interval than with perception of a control interval following a passive movement. We suggest that the supplementary motor complex contributes to the subjective experience of temporal flow that accompanies goal-directed voluntary actions.

Neuroimaging has demonstrated that the illusory self-attribution of body parts engages frontal and intraparietal brain areas, and recent evidence further suggests an involvement of visual body-selective regions in the occipitotemporal cortex. However, little is known about the principles of information exchange within this network. Here, using automated congruent versus incongruent visuotactile stimulation of distinct anatomical locations on the participant's right arm and a realistic dummy counterpart in an fMRI scanner, we induced an illusory self-attribution of the dummy arm. The illusion consistently activated a left-hemispheric network comprising ventral premotor cortex (PMv), intraparietal sulcus (IPS), and body-selective regions of the lateral occipitotemporal cortex (LOC). Importantly, during the illusion, the functional coupling of the PMv and the IPS with the LOC increased substantially, and dynamic causal modeling revealed a significant enhancement of connections from the LOC and the secondary somatosensory cortex to the IPS. These results comply with the idea that the brain's inference mechanisms rely on the hierarchical propagation of prediction error. During illusory self-attribution, unpredicted ambiguous sensory input about one's body configuration may result in the generation of such prediction errors in visual and somatosensory areas, which may be conveyed to parietal integrative areas.

61Stroke could increase body-ownership and agency over an external limb.61Stroke could limit changes in galvanic skin response and skin temperature.61Illusion of ownership could decrease electromyographic activity in stroke subjects.61It is hypothesized that the premotor cortex could cause these effects.61Results could evidence a body schema plasticity promoted by stroke.

Rubber-hand and virtual-hand illusions show that people can perceive body ownership for objects under suitable conditions. Bottom-up approaches assume that perceived ownership emerges from multisensory matching (e.g., between seen object and felt hand movements), whereas top-down approaches claim that novel body parts are integrated only if they resemble some part of a permanent internal body representation. We demonstrate that healthy adults perceive body ownership for a virtual balloon changing in size, and a virtual square changing in size or color, in synchrony with movements of their real hand. This finding is inconsistent with top-down approaches and amounts to an existence proof that non-corporeal events can be perceived as body parts if their changes are systematically related to one actions. It also implies that previous studies with passive-stimulation techniques might have underestimated the plasticity of body representations and put too much emphasis on the resemblance between viewed object and real hand.

Where are my hands? The brain can answer this question using sensory information arising from vision, proprioception, or touch. Other sources of information about the position of our hands can be derived from multisensory interactions (or potential interactions) with our close environment, such as when we grasp or avoid objects. The pioneering study of multisensory representations of peripersonal space was published in Behavioural Brain Research almost 30 years ago [Rizzolatti G, Scandolara C, Matelli M, Gentilucci M. Afferent properties of periarcuate neurons in macaque monkeys. II. Visual responses. Behav Brain Res 1981;2:147–63]. More recently, neurophysiological, neuroimaging, neuropsychological, and behavioural studies have contributed a wealth of evidence concerning hand-centred representations of objects in peripersonal space. This evidence is examined here in detail. In particular, we focus on the use of artificial dummy hands as powerful instruments to manipulate the brain's representation of hand position, peripersonal space, and of hand ownership. We also review recent studies of the ‘rubber hand illusion’ and related phenomena, such as the visual capture of touch, and the recalibration of hand position sense, and discuss their findings in the light of research on peripersonal space. Finally, we propose a simple model that situates the ‘rubber hand illusion’ in the neurophysiological framework of multisensory hand-centred representations of space.

AbstractConsiderable progress has been made in developing models of cerebellar function in sensorimotor control, as well as in identifying key problems that are the focus of current investigation. In this consensus paper, we discuss the literature on the role of the cerebellar circuitry in motor control, bringing together a range of different viewpoints. The following topics are covered: oculomotor control, classical conditioning (evidence in animals and in humans), cerebellar control of motor speech, control of grip forces, control of voluntary limb movements, timing, sensorimotor synchronization, control of corticomotor excitability, control of movement-related sensory data acquisition, cerebro-cerebellar interaction in visuokinesthetic perception of hand movement, functional neuroimaging studies, and magnetoencephalographic mapping of cortico-cerebellar dynamics. While the field has yet to reach a consensus on the precise role played by the cerebellum in movement control, the literature has witnessed the emergence of broad proposals that address cerebellar function at multiple levels of analysis. This paper highlights the diversity of current opinion, providing a framework for debate and discussion on the role of this quintessential vertebrate structure.

It is nearly 1002years since Patrick Haggard and colleagues first reported the ‘intentional binding’ effect ( Haggard, Clark, & Kalogeras, 2002 ). The intentional binding effect refers to the subjective compression of the temporal interval between a voluntary action and its external sensory consequence. Since the first report, considerable interest has been generated and a fascinating array of studies has accumulated. Much of the interest in intentional binding comes from the promise to shed light on human agency. In this review we survey studies on intentional binding, focusing, in particular, on the link between intentional binding and the sense of agency (the experience of controlling action to influence events in the environment). We suggest that, whilst it is yet to be fully explicated, the link between intentional binding and the sense of agency is compelling. We conclude by considering outstanding questions and future directions for research on intentional binding.

Recent advances in experimental science have made it possible to investigate the perceptual processes involved in generating a sense of owning an entire body. This is achieved by full-body ownership illusions which make use of specific patterns of visual and somatic stimuli integration. Here we investigate the fundamental question of the reference frames used in the process of attributing an entire body to the self. We quantified the strength of the body-swap illusion in conditions where the participants were observing this artificial body from the perspective of the first or third person. Consistent results from subjective reports and physiological recordings show that the first person visual perspective is critical for the induction of this full-body ownership illusion. This demonstrates that the multisensory integration processes producing the sense of corporeal self operates in an ego-centric reference frame.

Voluntary motor control over artificial hands has been shown to provoke a subjective incorporation of the artificial limb into body representations. However, in most studies projected or mirrored images of own hands were presented as ‘artificial’ body parts. Using the paradigm of the rubber hand illusion (RHI), we assessed the impact of tactile sensations and voluntary movements with respect to an unambiguously body-extraneous, artificial hand. In addition to phenomenal self-reports and pointing movements towards the own hand, we introduced a new procedure for perceptual judgements enabling the assessment of proprioceptive drift and judgement reliability regarding perceived hand location. RHI effects were comparable for tactile sensations and voluntary movements, but characteristic discrepancies were found for pointing movements. They were differently affected by the induction methods, and RHI effects were uncorrelated between both methods. These observations shed new light on inconsistent results concerning RHI effects on motor responses.

61Difficulty of action selection reduces the sense of agency over action outcomes.61Neural signals at the time of the action contribute to judgements of agency.61Neural processing of action outcomes also contributes to sense of agency.61Sense of agency integrates action- and outcome-related neural information.

Identifying with a body is central to being a conscious self. The now classic "rubber hand illusion" demonstrates that the experience of body-ownership can be modulated by manipulating the timing of exteroceptive (visual and tactile) body-related feedback. Moreover, the strength of this modulation is related to individual differences in sensitivity to internal bodily signals (interoception). However the interaction of exteroceptive and interoceptive signals in determining the experience of body-ownership within an individual remains poorly understood. Here, we demonstrate that this depends on the online integration of exteroceptive and interoceptive signals by implementing an innovative "cardiac rubber hand illusion" that combined computer-generated augmented-reality with feedback of interoceptive (cardiac) information. We show that both subjective and objective measures of virtual-hand ownership are enhanced by cardio-visual feedback in-time with the actual heartbeat, as compared to asynchronous feedback. We further show that these measures correlate with individual differences in interoceptive sensitivity, and are also modulated by the integration of proprioceptive signals instantiated using real-time visual remapping of finger movements to the virtual hand. Our results demonstrate that interoceptive signals directly influence the experience of body ownership via multisensory integration, and they lend support to models of conscious selfhood based on interoceptive predictive coding.

Each action has sensory consequences that need to be distinguished from sensations arising from the environment. This is accomplished by the comparing of internal predictions about these consequences with the actual afference, thereby isolating the afferent component that is self-produced. Because the sensory consequences of actions vary as a result of changes of the effector's efficacy, internal predictions need to be updated continuously and on a short time scale. Here, we tested the hypothesis that this updating of predictions about the sensory consequences of actions is mediated by the cerebellum, a notion that parallels the cerebellum's role in motor learning. Patients with cerebellar lesions and their matched controls were equally able to detect experimental modifications of visual feedback about their pointing movements. When such feedback was constantly rotated, both groups instantly attributed the visual feedback to their own actions. However, in interleaved trials without actual feedback, patients did no longer account for this feedback rotation--neither perceptually nor with respect to motor performance. Both deficits can be explained by an impaired updating of internal predictions about the sensory consequences of actions caused by cerebellar pathology. Thus, the cerebellum guarantees both precise performance and veridical perceptual interpretation of actions.

The neurocognitive structure of the acting self has recently been widely studied, yet is still perplexing and remains an often confounded issue in cognitive neuroscience, psychopathology and philosophy. We provide a new systematic account of two of its main features, the sense of agency and the sense of ownership, demonstrating that although both features appear as phenomenally uniform, they each in fact are complex crossmodal phenomena of largely heterogeneous functional and (self-)representational levels. These levels can be arranged within a gradually evolving, onto- and phylogenetically plausible framework which proceeds from basic non-conceptual sensorimotor processes to more complex conceptual and meta-representational processes of agency and ownership, respectively. In particular, three fundamental levels of agency and ownership processing have to be distinguished: The level of feeling, thinking and social interaction. This naturalistic account will not only allow to “ground the self in action”, but also provide an empirically testable taxonomy for cognitive neuroscience and a new tool for disentangling agency and ownership disturbances in psychopathology (e.g. alien hand, anarchic hand, anosognosia for one’s own hemiparesis).

Recent studies have shown that the well-known effect of multisensory stimulation on body-awareness can be extended to self-recognition. Seeing someone else's face being touched at the same time as one's own face elicits changes in the mental representation of the self-face. We sought to further elucidate the underlying mechanisms and the effects of interpersonal multisensory stimulation (IMS) on the mental representation of the self and others.Participants saw an unfamiliar face being touched synchronously or asynchronously with their own face, as if they were looking in the mirror. Following synchronous, but not asynchronous, IMS, participants assimilated features of the other's face in the mental representation of their own face as evidenced by the change in the point of subjective equality for morphed pictures of the two faces. Interestingly, synchronous IMS resulted in a unidirectional change in the self-other distinction, affecting recognition of one's own face, but not recognition of the other's face. The participants' autonomic responses to objects approaching the other's face were higher following synchronous than asynchronous IMS, but this increase was not specific to the pattern of IMS in interaction with the viewed object. Finally, synchronous, as compared to asynchronous, IMS resulted in significant differences in participants' ratings of their experience, but unlike other bodily illusions, positive changes in subjective experience were related to the perceived physical similarity between the two faces, and not to identification.Synchronous IMS produces quantifiable changes in the mental representations of one's face, as measured behaviorally. Changes in autonomic responses and in the subjective experience of self-identification were broadly consistent with patterns observed in other bodily illusions, but less robust. Overall, shared multisensory experiences between self and other can change the mental representation of one's identity, and the perceived similarity of others relative to one's self.

Empirical research on the bodily self has only recently started to investigate how the link between a body and the experience of this body as is developed, maintained or disturbed. The Rubber Hand Illusion has been used as a model instance of the normal sense of embodiment to investigate the processes that underpin the experience of body-ownership. This review puts forward a neurocognitive model according to which body-ownership arises as an interaction between current multisensory input and internal models of the body. First, a pre-existing stored model of the body distinguishes between objects that may or may not be part of one's body. Second, on-line anatomical and postural representations of the body modulate the integration of multisensory information that leads to the recalibration of visual and tactile coordinate systems. Third, the resulting referral of tactile sensation will give rise to the subjective experience of body-ownership. These processes involve a neural network comprised of the right temporoparietal junction which tests the incorporeability of the external object, the secondary somatosensory cortex which maintains an on-line representation of the body, the posterior parietal and ventral premotor cortices which code for the recalibration of the hand-centred coordinate systems, and the right posterior insula which underpins the subjective experience of body-ownership. The experience of body-ownership may represent a critical component of self-specificity as evidenced by the different ways in which multisensory integration in interaction with internal models of the body can actually manipulate important physical and psychological aspects of the self.

The experience of body ownership can be successfully manipulated during the rubber hand illusion using synchronous multisensory stimulation. The hypothesis that multisensory integration is both a necessary and sufficient condition for body ownership is debated. We systematically varied the appearance of the object that was stimulated in synchrony or asynchrony with the participant’s hand. A viewed object that was transformed in three stages from a plain wooden block to a wooden hand was compared to a realistic rubber hand. Introspective and behavioural results show that participants experience a sense of ownership only for the realistic prosthetic hand, suggesting that not all objects can be experienced as part of one’s body. Instead, the viewed object must fit with a reference model of the body that contains important structural information about body parts. This body model can distinguish between corporeal and non-corporeal objects, and it therefore plays a critical role in maintaining a coherent sense of one’s body.

Abstract Body ownership refers to the special perceptual status of one's own body, which makes bodily sensations seem unique to oneself. We studied the neural correlates of body ownership by controlling whether an external object was accepted as part of the body or not. In the rubber hand illusion (RHI), correlated visuotactile stimulation causes a fake hand to be perceived as part of one's own body. In the present study, we distinguished between the causes (i.e., multisensory stimulation) and the effect (i.e., the feeling of ownership) of the RHI. Participants watched a right or a left rubber hand being touched either synchronously or asynchronously with respect to their own unseen right hand. A quantifiable correlate of the RHI is a shift in the perceived position of the subject's hand toward the rubber hand. We used positron emission tomography to identify brain areas whose activity correlated with this proprioceptive measure of body ownership. Body ownership was related to activity in the right posterior insula and the right frontal operculum. Conversely, when the rubber hand was not attributed to the self, activity was observed in the contralateral parietal cortex, particularly the somatosensory cortex. These structures form a network that plays a fundamental role in linking current sensory stimuli to one's own body and thus also in self-consciousness.

Abstract Abstract: The comparator-model, originally developed to explain motor action, has recently been invoked to explain several aspects of the self. However, in the first place it may not be used to explain a basic self-world distinction because it presupposes one. Our alternative account is based on specific systematic covariation between action and perception. Secondly, the comparator model cannot explain the feeling of ownership of thoughts. We argue—contra Frith and Campbell—that thoughts are not motor processes and therefore cannot be described by the comparator-model. Rather, thoughts can be the triggering cause (intention) for actions. An alternative framework for the explanation of thought insertion in schizophrenics is presented.

Abstract A virtual-reality setup was used to investigate the relationship between perceived body ownership and subjective anxiety, as assessed by an anxiety inventory (SA-I). A pilot study confirmed that synchrony between the participant's real hand movements and the movements of a virtual effector induced perceived ownership illusions. The illusions were comparable for virtual human hands and virtual cat claws, even though the overall acceptance was greater for human hands. In Experiment 1, participants used the virtual effector to collect coins and avoid knives descending on a screen before anxiety was measured. The level of anxiety increased with synchrony and was higher for human hands than for cat claws, but these two effects were independent. Experiment 2 separated effects of coin catching and knife avoiding by means of a between-participant design. The outcome of Experiment 1 was replicated in the knife-avoiding task but not in the coin-catching task, in which anxiety levels were low and not systematically affected by the type of virtual effector. Taken altogether, our findings suggest that subjective anxiety and ownership are strongly related.

The tendency to perceive an artificial effector as part of one own body is known to depend on temporal criteria, like the synchrony between stimulus events informing about the effector. The role of spatial factors is less well understood. Rather than physical distance, which has been manipulated in previous studies, we investigated the role of relative, context-induced distance between the participant real hand and an artificial hand stimulated synchronously or asynchronously with the real hand. We replicated previously reported distance effects in a virtual reality setup: the perception of ownership increased with decreased distance, and the impact of synchrony was stronger for short distances. More importantly, we found that ownership perception and impact of synchrony were affected by previous distance: the same, medium distance between real and artificial hand induced more pronounced ownership after having experienced a far-distance condition than after a near-distance condition. This suggests that subjective, context-induced spatial reference frames contribute to ownership perception, which does not seem to fit with the idea of fixed spatial criteria and/or permanent body representations are the sole determinants of perceived body ownership.

We describe an illusion in which a stranger's voice, when presented as the auditory concomitant of a participant's own speech, is perceived as a modified version of their own voice. When the congruence between utterance and feedback breaks down, the illusion is also broken. Compared to a baseline condition in which participants heard their own voice as feedback, hearing a stranger's voice induced robust changes in the fundamental frequency (F0) of their production. Moreover, the shift in F0 appears to be feedback dependent, since shift patterns depended reliably on the relationship between the participant's own F0 and the stranger-voice F0. The shift in F0 was evident both when the illusion was present and after it was broken, suggesting that auditory feedback from production may be used separately for self-recognition and for vocal motor control. Our findings indicate that self-recognition of voices, like other body attributes, is malleable and context dependent.