The neuropsychology and neuroanatomy of reduplicative paramnesia.

Reduplicative paramnesia refers to the delusional belief that there are identical places in different locations. In this case-control study we investigated the clinical, phenomenological, neuropsychological and neuroanatomical data of eleven patients with reduplicative paramnesia and compared them against a control group of eleven patients with severe spatial disorientation without signs of reduplicative paramnesia. We show that most patients with reduplicative paramnesia report that a current place is reduplicated and/or relocated to an other familiar place. Patients with reduplicative paramnesia show a higher prevalence of deficits in the executive functions compared to the control patients, while mnestic and visuo-spatial deficits were both frequent in patients with reduplicative paramnesia and the control group. Patients with reduplicative paramnesia mostly suffer from right hemispheric lesions with a maximal overlap in the dorsolateral prefrontal cortex. Using lesion network mapping we show that lesions causing reduplicative paramnesia are connected to bilateral anterior insula and the right cingulate cortex. We argue that patients with reduplicative paramnesia fail to integrate the actual context with visuo-spatial memories and personal relevant emotional information due to a disruption of the neural network within the anterior temporal lobe, the cingulate cortex and the anterior insula. Also patients with reduplicative paramnesia are not able to resolve this conflict due to the lesion of the dorsolateral prefrontal cortex and executive dysfunction.

Augmented manipulation ability in humans with six-fingered hands.

Neurotechnology attempts to develop supernumerary limbs, but can the human brain deal with the complexity to control an extra limb and yield advantages from it? Here, we analyzed the neuromechanics and manipulation abilities of two polydactyly subjects who each possess six fingers on their hands. Anatomical MRI of the supernumerary finger (SF) revealed that it is actuated by extra muscles and nerves, and fMRI identified a distinct cortical representation of the SF. In both subjects, the SF was able to move independently from the other fingers. Polydactyly subjects were able to coordinate the SF with their other fingers for more complex movements than five fingered subjects, and so carry out with only one hand tasks normally requiring two hands. These results demonstrate that a body with significantly more degrees-of-freedom can be controlled by the human nervous system without causing motor deficits or impairments and can instead provide superior manipulation abilities.

Non-invasive brain stimulation of motor cortex induces embodiment when integrated with virtual reality feedback.

Previous evidence highlighted the multisensory-motor origin of embodiment - that is, the experience of having a body and of being in control of it - and the possibility of experimentally manipulating it. For instance, an illusory feeling of embodiment towards a fake hand can be triggered by providing synchronous visuo-tactile stimulation to the hand of participants and to a fake hand or by asking participants to move their hand and observe a fake hand moving accordingly (rubber hand illusion). Here, we tested whether it is possible to manipulate embodiment not through stimulation of the participant's hand, but by directly tapping into the brain's hand representation via non-invasive brain stimulation. To this aim, we combined transcranial magnetic stimulation (TMS), to activate the hand corticospinal representation, with virtual reality (VR), to provide matching (as contrasted to non-matching) visual feedback, mimicking involuntary hand movements evoked by TMS. We show that the illusory embodiment occurred when TMS pulses were temporally matched with VR feedback, but not when TMS was administered outside primary motor cortex, (over the vertex) or when stimulating motor cortex at a lower intensity (that did not activate peripheral muscles). Behavioural (questionnaires) and neurophysiological (motor-evoked-potentials, TMS-evoked-movements) measures further indicated that embodiment was not explained by stimulation per se, but depended on the temporal coherence between TMS-induced activation of hand corticospinal representation and the virtual bodily feedback. This reveals that non-invasive brain stimulation may replace the application of external tactile hand cues and motor components related to volition, planning and anticipation.

Changing motor perception by sensorimotor conflicts and body ownership.

Experimentally induced sensorimotor conflicts can result in a loss of the feeling of control over a movement (sense of agency). These findings are typically interpreted in terms of a forward model in which the predicted sensory consequences of the movement are compared with the observed sensory consequences. In the present study we investigated whether a mismatch between movements and their observed sensory consequences does not only result in a reduced feeling of agency, but may affect motor perception as well. Visual feedback of participants' finger movements was manipulated using virtual reality to be anatomically congruent or incongruent to the performed movement. Participants made a motor perception judgment (i.e. which finger did you move?) or a visual perceptual judgment (i.e. which finger did you see moving?). Subjective measures of agency and body ownership were also collected. Seeing movements that were visually incongruent to the performed movement resulted in a lower accuracy for motor perception judgments, but not visual perceptual judgments. This effect was modified by rotating the virtual hand (Exp.2), but not by passively induced movements (Exp.3). Hence, sensorimotor conflicts can modulate the perception of one's motor actions, causing viewed "alien actions" to be felt as one's own.

The cognitive and neural time course of empathy and sympathy: an electrical neuroimaging study on self-other interaction.

Although extensively investigated in socio-cognitive neuroscience, empathy is difficult to study. The first difficulty originates in its multifaceted nature. According to the multidimensional model, empathy combines emotional, automatic (simulation), cognitive (mentalizing) and regulatory (executive functions) processes. Substantial functional magnetic resonance imaging (fMRI) data demonstrated that co-activations in the mirror neuron system (MNS) and mentalizing network (MENT) sustain this co-recruitment of so-called first- and second-person-like processes. Because of the poor temporal resolution of fMRI techniques, we currently lack evidence about the precise timing of the MNS-MENT combination. An important challenge is, thus, to disentangle how MNS and MENT dynamically work together along time in empathy. Moreover, the role of the executive functions in the MNS-MENT combination time course is still unknown. Second, empathy - feeling into - is closely related to sympathy - feeling with - and both phenomena are often conflated in experimental studies on intersubjectivity. In this electrical neuroimaging (EEG) pilot-study, we tested whether the egocentered vs. heterocentered visuo-spatial mechanisms respectively associated with sympathy and empathy differentially modulate the dynamic combination of the MNS-MENT activations in their respective neural time course. For that, we employed our newly developed behavioral paradigm assessing the visuo-spatial - but not emotional - features of empathy and sympathy. Using a data-driven approach, we report that empathy and sympathy are underlied by sequential activations in the MNS from the insula to the inferior frontal gyrus (IFG) between 63ms and 424ms. However, at 333-424ms, empathy triggered greater co-activations in the right IFG and dorsolateral prefrontal cortex (dlPFC) (executive functions). Linking together our present and prior (Thirioux et al., 2010) findings from the same dataset, we suggest that this greater recruitment of the right dlPFC monitors the shift from egocentered and first-person-like mechanisms in the MNS to heterocentered and second-person-like mechanisms in the left temporo-parietal junction within the MENT, i.e., reflecting the onset of perspective-change processes in the neural time course of empathy. Contrasting with sympathy, this recruitment of the executive functions could modulate the output end of the mirroring processing in the premotor and sensorimotor cortices.

Self in motion: sensorimotor and cognitive mechanisms in gait agency.

Acting in our environment and experiencing ourselves as conscious agents are fundamental aspects of human selfhood. While large advances have been made with respect to understanding human sensorimotor control from an engineering approach, knowledge about its interaction with cognition and the conscious experience of movement (agency) is still sparse, especially for locomotion. We investigated these relationships by using life-size visual feedback of participants' ongoing locomotion, thereby extending agency research previously limited to goal-directed upper limb movements to continuous movements of the entire body. By introducing temporal delays and cognitive loading we were able to demonstrate distinct effects of bottom-up visuomotor conflicts as well as top-down cognitive loading on the conscious experience of locomotion (gait agency) and gait movements. While gait agency depended on the spatial and temporal congruency of the avatar feedback, gait movements were solely driven by its temporal characteristics as participants nonconsciously attempted to synchronize their gait with their avatar's gait. Furthermore, gait synchronization was suppressed by cognitive loading across all tested delays, whereas gait agency was only affected for selective temporal delays that depended on the participant's step cycle. Extending data from upper limb agency and auditory gait agency, our results are compatible with effector-independent and supramodal control of agency; they show that both mechanisms are dissociated from automated sensorimotor control and that cognitive loading further enhances this dissociation.

"Self pop-out": agency enhances self-recognition in visual search.

In real-life situations, we are often required to recognize our own movements among movements originating from other people. In social situations, these movements are often correlated (for example, when dancing or walking with others) adding considerable difficulty to self-recognition. Studies from visual search have shown that visual attention can selectively highlight specific features to make them more salient. Here, we used a novel visual search task employing virtual reality and motion tracking to test whether visual attention can use efferent information to enhance self-recognition of one's movements among four or six moving avatars. Active movements compared to passive movements allowed faster recognition of the avatar moving like the subject. Critically, search slopes were flat for the active condition but increased for passive movements, suggesting efficient search for active movements. In a second experiment, we tested the effects of using the participants' own movements temporally delayed as distractors in a self-recognition discrimination task. We replicated the results of the first experiment with more rapid self-recognition during active trials. Importantly, temporally delayed distractors increased reaction times despite being more perceptually different than the spatial distractors. The findings demonstrate the importance of agency in self-recognition and self-other discrimination from movement in social settings.

Visuo-tactile integration and body ownership during self-generated action.

Although there is increasing knowledge about how visual and tactile cues from the hands are integrated, little is known about how self-generated hand movements affect such multisensory integration. Visuo-tactile integration often occurs under highly dynamic conditions requiring sensorimotor updating. Here, we quantified visuo-tactile integration by measuring cross-modal congruency effects (CCEs) in different bimanual hand movement conditions with the use of a robotic platform. We found that classical CCEs also occurred during bimanual self-generated hand movements, and that such movements lowered the magnitude of visuo-tactile CCEs as compared to static conditions. Visuo-tactile integration, body ownership and the sense of agency were decreased by adding a temporal visuo-motor delay between hand movements and visual feedback. These data show that visual stimuli interfere less with the perception of tactile stimuli during movement than during static conditions, especially when decoupled from predictive motor information. The results suggest that current models of visuo-tactile integration need to be extended to account for multisensory integration in dynamic conditions.

I feel who I see: visual body identity affects visual-tactile integration in peripersonal space.

Recent studies have shown the importance of integrating multisensory information in the body representation for constituting self-consciousness. However, one idea that has received only scant attention is that our body representation is also constituted by knowledge of bodily visual characteristics (i.e. 'what I look like'). Here in two experiments we used a full body crossmodal congruency task in which visual distractors were presented on a photograph of the participant, another person, who was either familiar or unfamiliar, or an object. Results revealed that during the 'self-condition' CCEs were enhanced compared to the 'other condition'. The CCE was similar for unfamiliar and familiar others. CCEs for the object condition were significantly smaller. The results show that presentation of an irrelevant image of a body affects multimodal processing and that the effect is enhanced when that image is of the self. The results hold intriguing implications for body representation in social situations.

Early and late activity in somatosensory cortex reflects changes in bodily self-consciousness: an evoked potential study.

How can we investigate the brain mechanisms underlying self-consciousness? Recent behavioural studies on multisensory bodily perception have shown that multisensory conflicts can alter bodily self-consciousness such as in the "full body illusion" (FBI) in which changes in self-identification with a virtual body and tactile perception are induced. Here we investigated whether experimental changes in self-identification during the FBI are accompanied by activity changes in somatosensory cortex by recording somatosensory-evoked potentials (SEPs). To modulate self-identification, participants were filmed by a video camera from behind while their backs were stroked, either synchronously (illusion condition) or asynchronously (control condition) with respect to the stroking seen on their virtual body. Tibial nerve SEPs were recorded during the FBI and analysed using evoked potential (EP) mapping. Tactile mislocalisation was measured using the crossmodal congruency task. SEP mapping revealed five sequential periods of brain activation during the FBI, of which two differed between the illusion condition and the control condition. Activation at 30-50 ms (corresponding to the P40 component) in primary somatosensory cortex was stronger in the illusion condition. A later activation at ∼110-200 ms, likely originating in higher-tier somatosensory regions in parietal cortex, was stronger and lasted longer in the control condition. These data show that changes in bodily self-consciousness modulate activity in primary and higher-tier somatosensory cortex at two distinct processing steps. We argue that early modulations of primary somatosensory cortex may be a consequence of (1) multisensory integration of synchronous vs. asynchronous visuo-tactile stimuli and/or (2) differences in spatial attention (to near or far space) between the conditions. The later activation in higher-tier parietal cortex (and potentially other regions in temporo-parietal and frontal cortex) likely reflects the detection of visuo-tactile conflicts in the asynchronous condition.

The human vestibular cortex revealed by coordinate-based activation likelihood estimation meta-analysis.

The vestibular system contributes to the control of posture and eye movements and is also involved in various cognitive functions including spatial navigation and memory. These functions are subtended by projections to a vestibular cortex, whose exact location in the human brain is still a matter of debate (Lopez and Blanke, 2011). The vestibular cortex can be defined as the network of all cortical areas receiving inputs from the vestibular system, including areas where vestibular signals influence the processing of other sensory (e.g. somatosensory and visual) and motor signals. Previous neuroimaging studies used caloric vestibular stimulation (CVS), galvanic vestibular stimulation (GVS), and auditory stimulation (clicks and short-tone bursts) to activate the vestibular receptors and localize the vestibular cortex. However, these three methods differ regarding the receptors stimulated (otoliths, semicircular canals) and the concurrent activation of the tactile, thermal, nociceptive and auditory systems. To evaluate the convergence between these methods and provide a statistical analysis of the localization of the human vestibular cortex, we performed an activation likelihood estimation (ALE) meta-analysis of neuroimaging studies using CVS, GVS, and auditory stimuli. We analyzed a total of 352 activation foci reported in 16 studies carried out in a total of 192 healthy participants. The results reveal that the main regions activated by CVS, GVS, or auditory stimuli were located in the Sylvian fissure, insula, retroinsular cortex, fronto-parietal operculum, superior temporal gyrus, and cingulate cortex. Conjunction analysis indicated that regions showing convergence between two stimulation methods were located in the median (short gyrus III) and posterior (long gyrus IV) insula, parietal operculum and retroinsular cortex (Ri). The only area of convergence between all three methods of stimulation was located in Ri. The data indicate that Ri, parietal operculum and posterior insula are vestibular regions where afferents converge from otoliths and semicircular canals, and may thus be involved in the processing of signals informing about body rotations, translations and tilts. Results from the meta-analysis are in agreement with electrophysiological recordings in monkeys showing main vestibular projections in the transitional zone between Ri, the insular granular field (Ig), and SII.

Agency, gait and self-consciousness.

Agency is an important aspect of bodily self-consciousness, allowing us to separate own movements from those induced by the environment and to distinguish own movements from those of other agents. Unsurprisingly, theoretical frameworks for agency such as central monitoring are closely tied to computational models of sensorimotor control. Until recently agency research has largely focussed on goal-directed movements of the upper limbs. In particular, the influence of performance-related sensory cues and the relevance of prediction signals for agency judgements have been studied through a variety of spatio-temporal mismatches between movement and the sensory consequences of movement. However, agents often perform a different type of movement; highly automated movements that involve the entire body such as walking, cycling, and swimming with potentially different agency mechanisms. Here, we review recent work about agency for full-body movements such as gait, highlighting the effects of performance-related visual and auditory cues on gait agency. Gait movements differ from upper limb actions. Gait is cyclic, more rarely immediately goal-directed, and is generally considered one of the most automatic and unconscious actions. We discuss such movement differences with respect to the functional mechanisms of full-body agency and body-part agency by linking these gait agency paradigms to computational models of motor control. This is followed by a selective review of gait control, locomotion, and models of motor control relying on prediction signals and underlining their relevance for full-body agency.

Psychogenic amnesia and self-identity: a multimodal functional investigation.

BACKGROUND: Patients with psychogenic amnesia generally suffer from episodic memory deficits associated with an impairment of self-identity. While the first is generally attributed to limbic dysfunction, the latter might be related to posterior parietal cortex. METHODS AND RESULTS: In a patient with acute repetitive psychogenic amnesia, three different functional investigations (fMRI, electrical-neuroimaging, PET) during both resting-state and a behavioural paradigm testing 'time-travel' showed left posterior parietal activation, unlike in 12 control subjects. CONCLUSION: Impairment of self-identity and episodic memory in psychogenic amnesia may be associated with functional alterations of left posterior parietal cortex.

Spatiotemporal dynamics of visual vertical judgments: early and late brain mechanisms as revealed by high-density electrical neuroimaging.

Constructing and updating an internal model of verticality is fundamental for maintaining an erect posture and facilitating visuo-spatial processing. The judgment of the visual vertical (VV) has been intensively studied in psychophysical investigations and relies mainly on the integration of visual and vestibular signals, although a contribution of postural and somatosensory signals has been reported. Here we used high-density 192-channel evoked potential (EP) mapping and distributed source localization techniques to reveal the neural mechanisms of VV judgments. VV judgments (judging the orientation of visual lines with respect to the subjective vertical) were performed with and without a tilted visual frame. EP mapping revealed a sequence of neural processing steps (EP maps) of which two were specific for VV judgments. An early EP map, observed at ∼75-105 ms post-stimulus, was localized in right lateral temporo-occipital cortex. A later EP map (∼260-290 ms) was localized in bilateral temporo-occipital and parieto-occipital cortex. These data suggest that early VV-related neural processing involves the lateral and ventral visual stream and is related to visual processing concerning orientation, attention and comparison. The later, more dorsal, activation involves multimodal cortex subtending a constantly available and updated internal model of the vertical that we can refer to for the control of one's posture, actions, and visuo-spatial processing.

Neuroscience robotics to investigate multisensory integration and bodily awareness.

Humans experience the self as localized within their body. This aspect of bodily self-consciousness can be experimentally manipulated by exposing individuals to conflicting multisensory input, or can be abnormal following focal brain injury. Recent technological developments helped to unravel some of the mechanisms underlying multisensory integration and self-location, but the neural underpinnings are still under investigation, and the manual application of stimuli resulted in large variability difficult to control. This paper presents the development and evaluation of an MR-compatible stroking device capable of presenting moving tactile stimuli to both legs and the back of participants lying on a scanner bed while acquiring functional neuroimaging data. The platform consists of four independent stroking devices with a travel of 16-20 cm and a maximum stroking velocity of 15 cm/s, actuated over non-magnetic ultrasonic motors. Complemented with virtual reality, this setup provides a unique research platform allowing to investigate multisensory integration and its effects on self-location under well-controlled experimental conditions. The MR-compatibility of the system was evaluated in both a 3 and a 7 Tesla scanner and showed negligible interference with brain imaging. In a preliminary study using a prototype device with only one tactile stimulator, fMRI data acquired on 12 healthy participants showed visuo-tactile synchrony-related and body-specific modulations of the brain activity in bilateral temporoparietal cortex.

Carbon outclasses wood in racket paddles: Ratings of expert and intermediate tennis players.

Wooden racket paddles were modified with rubber and carbon fibre laminates and their differences tested in terms of flexural, damping, and coefficient of restitution properties. Four rackets types were designed: a wood reference, wood with rubber, carbon fibre 0°, and carbon fibre 90°. Seven expert and eight intermediate tennis players tested the rackets. To determine which of the four rackets suited the players best, we asked the players to compare the rackets two by two. After each pair tested, participants had to fill out a 4-item questionnaire in which different aspects of the rackets' performance were judged. The most preferred racket was the 0° carbon fibre racket, followed by the 90° carbon fibre racket, the wood racket and, finally, the 1-mm rubber racket. Thus, rackets with the highest stiffness, least damping, and highest coefficient of restitution were the most preferred. Interestingly, although experts and intermediate players overall judged the rackets in very similar ways according to force, vibration, and control, they were sensitive to quite different physical characteristics of the rackets.

The limits of agency in walking humans.

An important principle of human ethics is that individuals are not responsible for actions performed when unconscious. Recent research found that the generation of an action and the building of a conscious experience of that action (agency) are distinct processes and crucial mechanisms for self-consciousness. Yet, previous agency studies have focussed on actions of a finger or hand. Here, we investigate how agents consciously monitor actions of the entire body in space during locomotion. This was motivated by previous work revealing that (1) a fundamental aspect of self-consciousness concerns a single and coherent representation of the entire spatially situated body and (2) clinical instances of human behaviour without consciousness occur in rare neurological conditions such as sleepwalking or epileptic nocturnal wandering. Merging techniques from virtual reality, full-body tracking, and cognitive science of conscious action monitoring, we report experimental data about consciousness during locomotion in healthy participants. We find that agents consciously monitor the location of their entire body and its locomotion only with low precision and report that while precision remains low it can be systematically modulated in several experimental conditions. This shows that conscious action monitoring in locomoting agents can be studied in a fine-grained manner. We argue that the study of the mechanisms of agency for a person's full body may help to refine our scientific criteria of self-hood and discuss sleepwalking and related conditions as alterations in neural systems encoding motor awareness in walking humans.

Gender bending: auditory cues affect visual judgements of gender in biological motion displays.

The movement of an organism typically provides an observer with information in more than one sensory modality. The integration of information modalities reduces the likelihood that the observer will be confronted with a scene that is perceptually ambiguous. With that in mind, observers were presented with a series of point-light walkers each of which varied in the strength of the gender information they carried. Presenting those stimuli with auditory walking sequences containing ambiguous gender information had no effect on observers' ratings of visually perceived gender. When the visual stimuli were paired with auditory cues that were unambiguously female, observers' judgments of walker gender shifted such that ambiguous walkers were judged to look more female. To show that this is a perceptual rather than a cognitive effect, we induced visual gender after-effects with and without accompanying female auditory cues. The pairing of gender-neutral visual stimuli with unambiguous female auditory cues during adaptation elicited male after-effects. These data suggest that biological motion processing mechanisms can integrate auditory and visual cues to facilitate the extraction of higher-order features like gender. Possible neural substrates are discussed.

Three sequential brain activations encode mental transformations of upright and inverted human bodies: a high resolution evoked potential study.

Human bodies provide a particularly rich source of visual information. Whereas most previous studies have focused on the neural mechanisms during the perception and recognition of human bodies, the aim of the present study was to investigate the time course and location of brain activation during mental imagery of human bodies. When participants were asked to imagine themselves in the position of a visually presented human body as seen from many different angles and at two orientations (upright or inverted), their reaction times were faster for upright as compared to inverted bodies and correlated differently with the tested angles. These behavioral effects were also reflected in brain activation patterns, but only during the time period from 220 to 490 ms after stimulus onset. Evoked potential mapping and electrical neuroimaging revealed three distinct and sequential steps of processing related to mental body transformation: (1) an early activation in temporo-occipital and temporo-parietal cortex (220-360 ms) that does not distinguish between upright and inverted bodies, but closely reflects the effort of mental transformation, followed (2) by an activation in temporo-occipital and medial parieto-occipital cortex (350-460 ms) that encodes mental transformation for upright bodies, and (3) a later activation in temporo-occipital and prefrontal cortex (390-490 ms) that encodes mental transformation for inverted bodies. These data suggest that the mental transformation of human bodies is not a single process but a sequence of temporally distinct processing steps, where each step reflects a distinct aspect of the transformation process that consists of activations in a network of posterior brain areas including extrastriate cortex, temporo-parietal cortex, and medial parieto-occipital cortex, as well as an anterior brain region in prefrontal cortex.