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Abstract Self vs. external attribution of motions based on vestibular cues is suggested to underlie our coherent perception of object motion and self-motion. However, it remains unclear whether such attribution also underlies sensorimotor responses. Here, we examined this issue in the context of touch. We asked participants to lightly touch a moving object with their thumb while standing still on an unstable surface. We measured both the accuracy of judging the object motion direction and the postural response. If the attribution underlies both object-motion perception and posture control, sensitivity of posture to object motion should decrease with motion speed since high speed motion is unlikely to reflect self-motion. Furthermore, when motion perception is erroneous, there should be a corresponding increase in postural responses. Our results are consistent with these predictions and suggest that self-external attribution of somatosensory motion underlies both object motion perception and postural responses.

Abstract The sense of agency, the feeling of controlling one’s bodily actions and the world is altered in Depersonalisation (DP), a condition that makes people feel detached from one’s self and body. To investigate the link between depersonalisation and both implicit and explicit sense of agency, an online study was conducted using the influential Intentional Binding paradigm in a sample of non-clinical DP participants. The results did not reveal significant differences between individuals with low and high occurrences of DP experiences on the implicit and explicit sense of agency. However, participants with high occurrences of DP experiences showed a more time-sensitive explicit sense of agency and greater temporal distortions for short intervals in the absence of self-initiated motion. These results suggest that there is a discrepancy between implicit and explicit sense of agency in people with high levels of depersonalisation. Altogether, these findings call for further investigations of the key role of time perception on altered sense of self and agency in both non-clinical and clinical populations, to disentangle the mechanisms associated with the explicit and implicit sense of agency.

The law assumes that healthy adults are generally responsible for their actions and have the ability to control their behavior based on rational and moral principles. This contrasts with some recent neuroscientific accounts of action control. Nevertheless, both law and neuroscience acknowledge that strong emotions including fear and anger may “trigger” loss of normal voluntary control over action. Thus, “Loss of Control” is a partial defense for murder under English law, paralleling similar defenses in other legal systems. Here we consider the neuroscientific evidence for such legal classifications of responsibility, particularly focussing on how emotional states modulate voluntary motor control and sense of agency. First, we investigate whether neuroscience could contribute an evidence-base for law in this area. Second, we consider the societal impact of some areas where legal thinking regarding responsibility for action diverges from neuroscientific evidence: should we be guided by normative legal traditions, or by modern understanding of brain functions? In addressing these objectives, we propose a translation exercise between neuroscientific and legal terms, which may assist future interdisciplinary research.

Abstract Spatial acuity is a fundamental property of any sensory system. In the case of the somatosensory system, the two-point discrimination (2PD) test has long been used to investigate tactile spatial resolution. However, the somatosensory system comprises three main mechanoreceptive channels: the slowly adapting channel (SA) responds to steady pressure, the rapidly adapting channel (RA) responds to low-frequency vibration, and the Pacinian channel (PC) responds to high-frequency vibration. The use of mechanical stimuli in the classical 2PD test means that previous studies on tactile acuity have primarily focussed on the pressure-sensitive channel alone, while neglecting other submodalities. Here, we used a novel ultrasound stimulation to systematically investigate the spatial resolution of the two main vibrotactile channels. Contrary to the textbook view of poor spatial resolution for PC-like stimuli, across four experiments we found that high-frequency vibration produced surprisingly good spatial acuity. This effect remained after controlling for interchannel differences in stimulus detectability and perceived intensity. Laser doppler vibrometry experiments confirmed that the acuity of the PC channel was not simply an artifact of the skin’s resonance to high-frequency mechanical stimulation. Thus, PC receptors may transmit substantial spatial information, despite their sparse distribution, deep location, and large receptive fields.

Abstract The distributed nature of the neural substrate, and the difficulty of establishing necessity from correlative data, combine to render the mapping of brain function a far harder task than it seems. Methods capable of combining connective anatomical information with focal disruption of function are needed to disambiguate local from global neural dependence, and critical from merely coincidental activity. Here we present a comprehensive framework for focal and connective spatial inference based on sparse disruptive data, and demonstrate its application in the context of transient direct electrical stimulation of the human medial frontal wall during the pre-surgical evaluation of patients with focal epilepsy. Our framework formalizes voxel-wise mass-univariate inference on sparsely sampled data within the statistical parametric mapping framework, encompassing the analysis of distributed maps defined by any criterion of connectivity. Applied to the medial frontal wall, this transient dysconnectome approach reveals marked discrepancies between local and distributed associations of major categories of motor and sensory behaviour, revealing differentiation by remote connectivity to which purely local analysis is blind. Our framework enables disruptive mapping of the human brain based on sparsely sampled data with minimal spatial assumptions, good statistical efficiency, flexible model formulation, and explicit comparison of local and distributed effects.

Human–computer integration is an emerging area in which the boundary between humans and technology is blurred as users and computers work collaboratively and share agency to execute tasks. The sense of agency (SoA) is an experience that arises by a combination of a voluntary motor action and sensory evidence whether the corresponding body movements have somehow influenced the course of external events. The SoA is not only a key part of our experiences in daily life but also in our interaction with technology as it gives us the feeling of “I did that” as opposed to “the system did that,” thus supporting a feeling of being in control. This feeling becomes critical with human–computer integration, wherein emerging technology directly influences people’s body, their actions, and the resulting outcomes. In this review, we analyse and classify current integration technologies based on what we currently know about agency in the literature, and propose a distinction between body augmentation, action augmentation, and outcome augmentation. For each category, we describe agency considerations and markers of differentiation that illustrate a relationship between assistance level (low, high), agency delegation (human, technology), and integration type (fusion, symbiosis). We conclude with a reflection on the opportunities and challenges of integrating humans with computers, and finalise with an expanded definition of human–computer integration including agency aspects which we consider to be particularly relevant. The aim this review is to provide researchers and practitioners with guidelines to situate their work within the integration research agenda and consider the implications of any technologies on SoA, and thus overall user experience when designing future technology.

In this study, the authors distinguish between changes of mind about perceptual vs. intentional decisions. A Hierarchical Attractor Network Model is proposed in which human voluntary actions emerge from continuous and dynamic integration of higher-order intentions with sensory evidence and motor costs.

Working in military structures implies a reduction in individual autonomy, in which agents must comply with hierarchical orders. Here, the authors show that working within such a structure is associated with a reduced sense of agency and outcome processing for junior cadets, but this relationship is absent in trained officers.

The paper discusses the contribution that the neuroscience of action can offer to the legal understanding of action control and responsibility in the case of adult individuals. In particular, we address the issues that follow. What are the cognitive capacities that agents must display in order to be held liable to punishment in criminal law? Is the legal model of liability to punishment compatible with a scientifically informed understanding of voluntary behaviour? To what extent should the law take into account people’s subjective feelings about their own actions? As a result of our analyses, we indicate some areas where the contribution of the neuroscience of action to the law is potentially relevant. We focus on the subjectivity mechanisms of action control, specifically the requirement that the agent must violate the law voluntarily in order to be held responsible, and on the factors that modulate the wrongdoer’s experience of agency. Overall, we advocate more cross-disciplinary work, aimed to bridge the gap between conceptual boundaries, on the theme of responsibility for actions.

To interact meaningfully with its environment, an agent must integrate external information with its own internal states. However, information about the environment is often noisy. In this study, we identify a neural correlate that tracks how asymmetries between competing alternatives evolve over the course of a decision. In our task participants had to monitor a stream of discrete visual stimuli over time and decide whether or not to act, on the basis of either strong or ambiguous evidence. We found that the classic P3 event-related potential evoked by sequential evidence items tracked decision-making processes and predicted participants’ categorical choices on a single trial level, both when evidence was strong and when it was ambiguous. The P3 amplitudes in response to evidence supporting the eventually selected option increased over trial time as decisions evolved, being maximally different from the P3 amplitudes evoked by competing evidence at the time of decision. Computational modelling showed that both the neural dynamics and behavioural primacy and recency effects can be explained by a combination of (a) competition between mutually inhibiting accumulators for the two categorical choice outcomes, and (b) a context-dependant urgency signal. In conditions where evidence was presented at a low rate, urgency increased faster than in conditions when evidence was very frequent. We also found that the readiness potential, a classic marker of endogenously initiated actions, was observed preceding movements in all conditions - even when those were strongly driven by external evidence.

Milgram’s classical studies famously suggested a widespread willingness to obey authority, even to the point of inflicting harm. Important situational factors supporting obedience, such as proximity with the victim, have been established. Relatively little work has focused on how coercion affects individual cognition, or on identifying the cognitive factors that underlie inter-individual differences in the tendency to yield to coercion. Here, we used fMRI to investigate the neural systems associated with changes in volitional processes associated with sense of agency and sense of responsibility under coercion. Participants either freely chose, or were instructed by the experimenter, to give mildly painful electric shocks to another participant, or to refrain from doing so. We have previously shown that coercion reduces temporal binding, which has been proposed as an implicit proxy measure of sense of agency. We tested how reduced agency under coercion related to differences in neural activity between free choice and coercion. In contrast to previous studies and to participants performing the task outside the MRI scanner, on average there was no effect of coercion on agency for participants in the scanner. However, greater activity in the medial frontal gyrus was reliably associated with greater agency under coercion. A similar association was found using explicit responsibility ratings. Our findings suggest that medial frontal processes, perhaps related to volition during action planning and execution, may help to preserve a sense of accountability under coercion. Further, participants who administered more shocks under free choice showed reduced activity during free choice trials in brain areas associated with social cognition. Possibly, this might reflect participants cognitively distancing themselves from the recipient of the shocks under free choice, whereas this was not observed under coercion.

Attention plays a crucial role in functional neurological disorders. Attention to the symptoms leads to their exacerbation and distraction to their improvement or even transitory disappearance.Objective: The aim was to test if the alerting, orienting and particularly the executive aspect of attention are affected in functional movement disorders.Methods: Thirty patients with a functional movement disorder, 30 patients with an organic movement disorder and 30 healthy controls performed the attention network test.Results: The alerting and orienting effects were normal, but executive control of attention under conflict was abnormal in patients with functional movement disorders, compared to patients with an organic movement disorder and healthy controls.Conclusion: Executive dysfunction seems to be an important secondary feature of functional movement disorders, due to the overutilization of attentional resources for explicit movement control. Furthermore, it provides an explanation for seemingly unrelated symptoms commonly associated with functional movement disorders, such as concentration difficulties and fatigue.

Background: In functional movement disorders, explicit movements are impaired, while implicit movements are preserved. Furthermore, there is evidence that the sense of agency is abnormal.Aim: We aimed to investigate how motor responses and sense of agency were affected by subliminal or supraliminal cues in people with functional movement disorders.Methods: Twenty-three people with a functional movement disorder and 26 healthy controls took part in a subliminal and supraliminal priming experiment which investigated reaction times, choice and sense of agency. Participants pressed a left or right arrow key in response to an imperative left or right pointing arrow. Either key could be pressed in response to bidirectional arrows. The imperative arrow was preceded by a small left or right pointing prime arrow, that was non-predictive (50% correct) and was presented in either subliminal or supraliminal conditions. The participant's response caused the appearance of a colored circle and they rated the degree of control they felt over its appearance (sense of agency). The circle's color depended on whether their response was congruent or incongruent with the prime arrow direction. After exclusion, 19 participants remained in each group.Results: Prime-compatible responses led to faster reaction times in both the subliminal and supraliminal condition. Subliminal prime-compatible responses were chosen more frequently in the free choice condition. The sense of agency did not depend on prime-response congruency. There were no significant differences in any of these measures between the two groups.Conclusion: With non-predictive cues, reaction times, choices, and the sense of agency remain normal in people with functional movement disorders, for both subliminal and supraliminal primes. The findings suggest that it is not so much conscious awareness of the movement, but rather conscious motor preparation that is detrimental to motor function in functional movement disorders.

The Readiness Potential (RP) is a slow negative EEG potential found in the seconds preceding voluntary actions. Here, we explore whether the RP is found only at this time, or if it also occurs when no action is produced. Recent theories suggest the RP reflects the average of accumulated stochastic fluctuations in neural activity, rather than a specific signal related to self-initiated action: RP-like events should then be widely present, even in the absence of actions. We investigated this hypothesis by searching for RP-like events in background EEG of an appropriate dataset for which the action-locked EEG had previously been analysed to test other hypotheses [Khalighinejad, N., Brann, E., Dorgham, A., Haggard, P. Dissociating cognitive and motoric precursors of human self-initiated action. Journal of Cognitive Neuroscience. 2019, 1-14]. We used the actual mean RP as a template, and searched the entire epoch for similar neural signals, using similarity metrics that capture the temporal or spatial properties of the RP. Most EEG epochs contained a number of events that were similar to the true RP, but did not lead directly to any voluntary action. However, these RP-like events were equally common in epochs that eventually terminated in voluntary actions as in those where voluntary actions were not permitted. Events matching the temporal profile of the RP were also a poor match for the spatial profile, and vice versa. We conclude that these events are false positives, and do not reflect the same mechanism as the RP itself. Finally, applying the same template-search algorithm to simulated EEG data synthesized from different noise distributions showed that RP-like events will occur in any dataset containing the 1⁄f noise ubiquitous in EEG recordings. To summarise, we found no evidence of genuinely RP-like events at any time other than immediately prior to self-initiated actions. Our findings do not support a purely stochastic model of RP generation, and suggest that the RP may be a specific precursor of self-initiated voluntary actions.

Abstract Sense of agency (SoA), a feeling that one’s voluntary actions produce events in the external world, is a key factor behind every goal-directed human behaviour. Recent studies have demonstrated that SoA is reduced when one’s voluntary action causes negative outcomes, compared to when it causes positive outcomes. It is yet unclear whether this emotional modulation of SoA is caused by predicting the outcome valence (prediction hypothesis) or by retrospectively interpreting the outcome (postdiction hypothesis). To address this, we emulated a social situation where one’s voluntary action was followed by either another’s negative emotional vocalisation or positive emotional vocalisation. Crucially, the relation between an action and the emotional valence of its outcome was predictable in some blocks of trials, but unpredictable in other blocks. Quantitative, implicit measures of SoA based on the intentional binding effect supported the prediction hypothesis. Our findings imply that the social-emotional modulation of SoA is based on predicting the emotional valence of action outcomes.

There are few concepts as relevant as that of intentional action in shaping our sense of self and the interaction with the environment. At the same time, few concepts are so elusive. Indeed, both conceptual and neuroscientific accounts of intentional agency have proven to be problematic. On the one hand, most conceptual views struggle in defining how agents can adequately exert control over their actions. On the other hand, neuroscience settles for definitions by exclusion whereby key features of human intentional actions, including goal-directness, remain underspecified. This paper reviews the existing literature and sketches how this gap might be filled. In particular, we defend a gradualist notion of intentional behavior, which revolves around the following key features: autonomy, flexibility in the integration of causal vectors, and control.

The Kohnstamm phenomenon is a prolonged involuntary aftercontraction following a sustained voluntary isometric muscle contraction. The control principles of the Kohnstamm have been investigated using mechanical perturbations, but previous studies could not dissociate sensorimotor responses to perturbation from effects of gravity. We induced a horizontal, gravity-independent Kohnstamm movement around the shoulder joint, and applied resistive or assistive torques of 0.5 Nm after 20° angular displacement. A No perturbation control condition was included. Further, participants made velocity-matched voluntary movements, with or without similar perturbations, yielding a 2 × 3 factorial design. Resistive perturbations produced an increase in agonist electromyography (EMG), in both Kohnstamm and voluntary movements, while assistive perturbations produced a decrease. While overall Kohnstamm EMGs were greater than voluntary EMGs, the EMG responses to perturbation, when expressed as a percentage of unperturbed EMG activity, were significantly smaller during Kohnstamm movements than during voluntary movements. The results suggest that the Kohnstamm aftercontraction involves a central drive, coupled with low-gain servo control by a negative feedback loop between afferent input and a central motor command. The combination of strong efferent drive with low reflex gain may characterize involuntary control of postural muscles. Our results question traditional accounts involving purely reflexive mechanisms of postural maintenance. They also question existing high-gain, peripheral accounts of the Kohnstamm phenomenon, as well as accounts involving a central adaptation interacting with muscle receptors via a positive force feedback loop.

In human societies, agents are assumed to experience being the author of their own actions. These basic motoric experiences of action are influenced by social hierarchies, leading to surprising and morally significant results. Here we ask whether, under coercion, the sense of agency and responsibility pass from the person who receives orders to the person who gives them. Volunteers took turns to play the roles of 'commander', 'agent' or 'victim' in a task where the commander coerced the agent to deliver painful shocks to the 'victim'. We used 'intentional binding' as an implicit measure of sense of agency in both commanders and agents, in conditions of coercion and free-choice. We observed a reduced sense of agency when agents received coercive instructions, relative to when they freely chose which action to execute. We also found that sense of agency in the commanders was reduced when they coerced agents to administer the shock on their behalf, relative to when they acted by themselves. This last effect was associated with the commander's self-reported level on a psychopathy scale. Thus, coercion resulted in neither commander nor agent feeling agency for the effect of the action, as measured through implicit methods. Our results could have profound implications for social decision-making and social regulation of moral behaviour.

Perceptual decisions are classically thought to depend mainly on the stimulus characteristics, probability and associated reward. However, in many cases, the motor response is considered to be a neutral output channel that only reflects the upstream decision. Contrary to this view, we show that perceptual decisions can be recursively influenced by the physical resistance applied to the response. When participants reported the direction of the visual motion by left or right manual reaching movement with different resistances, their reports were biased towards the direction associated with less effortful option. Repeated exposure to such resistance on hand during perceptual judgements also biased subsequent judgements using voice, indicating that effector-dependent motor costs not only biases the report at the stage of motor response, but also changed how the sensory inputs are transformed into decisions. This demonstrates that the cost to act can influence our decisions beyond the context of the specific action.

Whether a visual stimulus seems near or far away depends partly on its vertical elevation. Contrasting theories suggest either that perception of distance could vary with elevation, because of memory of previous upwards efforts in climbing to overcome gravity, or because of fear of falling associated with the downwards direction. The vestibular system provides a fundamental signal for the downward direction of gravity, but the relation between this signal and depth perception remains unexplored. Here we report an experiment on vestibular contributions to depth perception, using Virtual Reality. We asked participants to judge the absolute distance of an object presented on a plane at different elevations during brief artificial vestibular inputs. Relative to distance estimates collected with the object at the level of horizon, participants tended to overestimate distances when the object was presented above the level of horizon and the head was tilted upward and underestimate them when the object was presented below the level of horizon. Interestingly, adding artificial vestibular inputs strengthened these distance biases, showing that online multisensory signals, and not only stored information, contribute to such distance illusions. Our results support the gravity theory of depth perception, and show that vestibular signals make an on-line contribution to the perception of effort, and thus of distance.

The sense of controlling one’s own actions is fundamental to normal human mental function, and also underlies concepts of social responsibility for action. However, it remains unclear how the wider social context of human action influences sense of agency. Using a simple experimental design, we investigated, for the first time, how observing the action of another person or a robot could potentially influence one’s own sense of agency. We assessed how observing another’s action might change the perceived temporal relationship between one’s own voluntary actions and their outcomes, which has been proposed as an implicit measure of sense of agency. Working in pairs, participants chose between two action alternatives, one rewarded more frequently than the other, while watching a rotating clock hand. They judged, in separate blocks, either the time of their own action, or the time of a tone that followed the action. These were compared to baseline judgements of actions alone, or tones alone, to calculate the perceptual shift of action toward outcome and vice versa. Our design focused on how these two dependent variables, which jointly provide an implicit measure of sense of agency, might be influenced by observing another’s action. In the observational group, each participant could see the other’s actions. Multivariate analysis showed that the perceived time of action and tone shifted progressively toward the actual time of outcome with repeated experience of this social situation. No such progressive change occurred in other groups for whom a barrier hid participants’ actions from each other. However, a similar effect was observed in the group that viewed movements of a human-like robotic hand, rather than actions of another person. This finding suggests that observing the actions of others increases the salience of the external outcomes of action and this effect is not unique to observing human agents. Social contexts in which we see others controlling external events may play an important role in mentally representing the impact of our own actions on the external world.

An Asian spice, Szechuan pepper (sanshool), is well known for the tingling sensation it induces on the mouth and on the lips. Electrophysiological studies have revealed that its active ingredient can induce firing of mechanoreceptor fibres that typically respond to mechanical vibration. Moreover, a human behavioral study has reported that the perceived frequency of sanshool-induced tingling matches with the preferred frequency range of the tactile rapidly adapting (RA) channel, suggesting the contribution of sanshool-induced RA channel firing to its unique perceptual experience. However, since the RA channel may not be the only channel activated by sanshool, there could be a possibility that the sanshool tingling percept may be caused in whole or in part by other sensory channels. Here, by using a perceptual interference paradigm, we show that the sanshool-induced RA input indeed contributes to the human tactile processing. The absolute detection thresholds for vibrotactile input were measured with and without sanshool application on the fingertip. Sanshool significantly impaired detection of vibrations at 30 Hz (RA channel dominant frequency), but did not impair detection of higher frequency vibrations at 240 Hz (Pacinian-corpuscle (PC) channel dominant frequency) or lower frequency vibrations at 1 Hz (slowly adapting 1 (SA1) channel dominant frequency). These results show that the sanshool induces a peripheral RA channel activation that is relevant for tactile perception. This anomalous activation of RA channels may contribute to the unique tingling experience of sanshool.

Does sense of agency (SoA) arise merely from action-outcome associations, or does an additional real-time process track each step along the chain? Tracking control predicts that deviant intermediate steps between action and outcome should reduce SoA. In two experiments, participants learned mappings between two finger actions and two tones. In later test blocks, actions triggered a robot hand moving either the same or a different finger, and also triggered tones, which were congruent or incongruent with the mapping. The perceived delay between actions and tones gave a proxy measure for SoA. Action-tone binding was stronger for congruent than incongruent tones, but only when the robot movement was also congruent. Congruent tones also had reduced N1 amplitudes, but again only when the robot movement was congruent. We suggest that SoA partly depends on a real-time tracking control mechanism, since deviant intermediate action of the robot reduced SoA over the tone.

Vestibular inputs are constantly processed and integrated with signals from other sensory modalities, such as vision and touch. The multiply-connected nature of vestibular cortical anatomy led us to investigate whether vestibular signals could participate in a multi-way interaction with visual and somatosensory perception. We used signal detection methods to identify whether vestibular stimulation might interact with both visual and somatosensory events in a detection task. Participants were instructed to detect near-threshold somatosensory stimuli that were delivered to the left index finger in one half of experimental trials. A visual signal occurred close to the finger in half of the trials, independent of somatosensory stimuli. A novel Near infrared caloric vestibular stimulus (NirCVS) was used to artificially activate the vestibular organs. Sham stimulations were used to control for non-specific effects of NirCVS. We found that both visual and vestibular events increased somatosensory sensitivity. Critically, we found no evidence for supra-additive multisensory enhancement when both visual and vestibular signals were administered together: in fact, we found a trend towards sub-additive interaction. The results are compatible with a vestibular role in somatosensory gain regulation.

The ability to decode an individual's intentions in real time has long been a 'holy grail' of research on human volition. For example, a reliable method could be used to improve scientific study of voluntary action by allowing external probe stimuli to be delivered at different moments during development of intention and action. Several Brain Computer Interface applications have used motor imagery of repetitive actions to achieve this goal. These systems are relatively successful, but only if the intention is sustained over a period of several seconds; much longer than the timescales identified in psychophysiological studies for normal preparation for voluntary action. We have used a combination of sensorimotor rhythms and motor imagery training to decode intentions in a single-trial cued-response paradigm similar to those used in human and non-human primate motor control research. Decoding accuracy of over 0.83 was achieved with twelve participants. With this approach, we could decode intentions to move the left or right hand at sub-second timescales, both for instructed choices instructed by an external stimulus and for free choices generated intentionally by the participant. The implications for volition are considered.

Vestibular signals are strongly integrated with information from several other sensory modalities. For example, vestibular stimulation was reported to improve tactile detection. However, this improvement could reflect either a multimodal interaction or an indirect interaction driven by vestibular effects on spatial attention and orienting. Here we investigate whether natural vestibular activation induced by passive whole-body rotation influences tactile detection. In particular, we assessed the ability to detect faint tactile stimuli to the fingertips of the left and right hand during spatially congruent or incongruent rotations. We found that passive whole-body rotations significantly enhanced sensitivity to faint shocks, without affecting response bias. Critically, this enhancement of somatosensory sensitivity did not depend on the spatial congruency between the direction of rotation and the hand stimulated. Thus, our results support a multimodal interaction, likely in brain areas receiving both vestibular and somatosensory signals.

Despite extensive research on face perception, few studies have investigated individuals' knowledge about the physical features of their own face. In this study, 50 participants indicated the location of key features of their own face, relative to an anchor point corresponding to the tip of the nose, and the results were compared to the true location of the same individual's features from a standardised photograph. Horizontal and vertical errors were analysed separately. An overall bias to underestimate vertical distances revealed a distorted face representation, with reduced face height. Factor analyses were used to identify separable subconfigurations of facial features with correlated localisation errors. Independent representations of upper and lower facial features emerged from the data pattern. The major source of variation across individuals was in representation of face shape, with a spectrum from tall/thin to short/wide representation. Visual identification of one's own face is excellent, and facial features are routinely used for establishing personal identity. However, our results show that spatial knowledge of one's own face is remarkably poor, suggesting that face representation may not contribute strongly to self-awareness.

Inhibition of prepotent action is an important aspect of self-control, particularly in social contexts. Action inhibition and its neural bases have been extensively studied. However, the neural precursors of free decisions to inhibit have hardly been studied. We asked participants to freely choose to either make a rapid key press in response to a visual cue, or to transiently inhibit action, and briefly delay responding. The task required a behavioural response on each trial, so trials involving inhibition could be distinguished from those without inhibition as those showing slower reaction times. We used this criterion to classify free-choice trials as either rapid or inhibited/delayed. For 13 participants, we measured the mean amplitude of the ERP activity at electrode Cz in three subsequent 50 ms time windows prior to the onset of the signal that either instructed to respond or inhibit, or gave participants a free choice. In two of these 50 ms time windows (-150 to -100, and -100 to -50 ms relative to action onset), the amplitude of prestimulus ERP differed between trials where participants "freely" chose whether to inhibit or to respond rapidly. Larger prestimulus ERP amplitudes were associated with trials in which participants decided to act rapidly as compared to trials in which they decided to delay their responses. Last-moment decisions to inhibit or delay may depend on unconscious preparatory neural activity.

Previous studies have shown that the perceived times of voluntary actions and their effects are perceived as shifted towards each other, so that the interval between action and outcome seems shortened. This has been referred to as 'intentional binding' (IB). However, the generality of this effect remains unclear. Here we demonstrate that Intentional Binding also occurs in complex control situations. Using an aircraft supervision task with different autopilot settings, our results first indicated a strong relation between measures of IB and different levels of system automation. Second, measures of IB were related to explicit agency judgement in this applied setting. We discuss the implications for the underlying mechanisms, and for sense of agency in automated environments.

Sense of agency refers to the feeling that "I" am responsible for those external events that are directly produced by one's own voluntary actions. Recent theories distinguish between a non-conceptual "feeling" of agency linked to changes in the processing of self-generated sensory events, and a higher-order judgement of agency, which attributes sensory events to the self. In the current study we explore the neural correlates of the judgement of agency by means of electrophysiology. We measured event-related potentials to tones that were either perceived or not perceived as triggered by participants' voluntary actions and related these potentials to later judgements of agency over the tones. Replicating earlier findings on predictive sensory attenuation, we found that the N1 component was attenuated for congruent tones that corresponded to the learned action-effect mapping as opposed to incongruent tones that did not correspond to the previously acquired associations between actions and tones. The P3a component, but not the N1, directly reflected the judgement of agency: deflections in this component were greater for tones judged as self-generated than for tones judged as externally produced. The fact that the outcome of the later agency judgement was predictable based on the P3a component demonstrates that agency judgements incorporate early information processing components and are not purely reconstructive, post-hoc evaluations generated at time of judgement.

BackgroundRepresenting one's own body is often viewed as a basic form of self-awareness. However, little is known about structural representations of the body in the brain.Methods and findingsWe developed an inter-manual version of the classical "in-between" finger gnosis task: participants judged whether the number of untouched fingers between two touched fingers was the same on both hands, or different. We thereby dissociated structural knowledge about fingers, specifying their order and relative position within a hand, from tactile sensory codes. Judgments following stimulation on homologous fingers were consistently more accurate than trials with no or partial homology. Further experiments showed that structural representations are more enduring than purely sensory codes, are used even when number of fingers is irrelevant to the task, and moreover involve an allocentric representation of finger order, independent of hand posture.ConclusionsOur results suggest the existence of an allocentric representation of body structure at higher stages of the somatosensory processing pathway, in addition to primary sensory representation.

In performing arts, body postures are both means for expressing an artist's intentions, and also artistic objects, appealing to the audience. The postures of classical ballet obey the body's biomechanical limits, but also follow strict rules established by tradition. This combination offers a perfect milieu for assessing scientifically how the execution of this particular artistic activity has changed over time, and evaluating what factors may induce such changes. We quantified angles between body segments in archive material showing dancers from a leading company over a 60-year period. The data showed that body positions supposedly fixed by codified choreography were in fact implemented by very different elevation angles, according to the year of ballet production. Progressive changes lead to increasingly vertical positions of the dancer's body over the period studied. Experimental data showed that these change reflected aesthetic choices of naïve modern observers. Even when reduced to stick figures and unrecognisable shapes, the more vertical postures drawn from later productions were systematically preferred to less vertical postures from earlier productions. This gradual change within a conservative art form provides scientific evidence that aesthetic change may arise from continuous interaction between artistic tradition, individual artists' creativity, and a wider environmental context. This context may include social aesthetic pressure from audiences.

When a part of the body moves, the sensation evoked by a probe stimulus to that body part is attenuated. Two mechanisms have been proposed to explain this robust and general effect. First, feedforward motor signals may modulate activity evoked by incoming sensory signals. Second, reafferent sensation from body movements may mask the stimulus. Here we delivered probe stimuli to the right index finger just before a cue which instructed subjects to make left or right index finger movements. When left and right cues were equiprobable, we found attenuation for stimuli to the right index finger just before this finger was cued (and subsequently moved). However, there was no attenuation in the right finger just before the left finger was cued. This result suggests that the movement made in response to the cue caused 'postdictive' attenuation of a sensation occurring prior to the cue. In a second experiment, the right cue was more frequent than the left. We now found attenuation in the right index finger even when the left finger was cued and moved. This attenuation linked to a movement that was likely but did not in fact occur, suggests a new expectation-based mechanism, distinct from both feedforward motor signals and postdiction. Our results suggest a new mechanism in motor-sensory interactions in which the motor system tunes the sensory inputs based on expectations about future possible actions that may not, in fact, be implemented.