a b s t r a c tAn 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 selfhood and discuss sleepwalking and related conditions as alterations in neural systems encoding motor awareness in walking humans.

Voluntary action is a fundamental element of self-consciousness. The readiness potential (RP), a slow drift of neural activity preceding self-initiated movement, has been suggested to reflect neural processes underlying the preparation of voluntary action; yet more than fifty years after its introduction, interpretation of the RP remains controversial. Based on previous research showing that internal bodily signals affect sensory processing and ongoing neural activity, we here investigated the potential role of interoceptive signals in voluntary action and the RP. We report that (1) participants initiate voluntary actions more frequently during expiration, (2) this respiration-action coupling is absent during externally triggered actions, and (3) the RP amplitude is modulated depending on the respiratory phase. Our findings demonstrate that voluntary action is coupled with the respiratory system and further suggest that the RP is associated with fluctuations of ongoing neural activity that are driven by the involuntary and cyclic motor act of breathing.

Kannape OA, Blanke O. Self in motion: sensorimotor and cognitive mechanisms in gait agency. J Neurophysiol 110: 1837-1847, 2013. First published July 3, 2013 doi:10.1152/jn.01042.2012.-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.

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.

The seemingly stable construct of our bodily self depends on the continued, successful integration of multisensory feedback about our body, rather than its purely physical composition. Accordingly, pathological disruption of such neural processing is linked to striking alterations of the bodily self, ranging from limb misidentification to disownership, and even the desire to amputate a healthy limb. While previous embodiment research has relied on experimental setups using supernumerary limbs in variants of the Rubber Hand Illusion, we here used Mixed Reality to directly manipulate the feeling of ownership for one's own, biological limb. Using a Head-Mounted Display, participants received visual feedback about their own arm, from an embodied first-person perspective. In a series of three studies, in independent cohorts, we altered embodiment by providing visuotactile feedback that could be synchronous (control condition) or asynchronous (400ms delay, Real Hand Illusion). During the illusion, participants reported a significant decrease in ownership of their own limb, along with a lowered sense of agency. Supporting the right-parietal body network, we found an increased illusion strength for the left upper limb as well as a modulation of the feeling of ownership during anodal transcranial direct current stimulation. Extending previous research, these findings demonstrate that a controlled, visuotactile conflict about one's own limb can be used to directly and systematically modulate ownershipwithout a proxy. This not only corroborates the malleability of body representation but questions its permanence. These findings warrant further exploration of combined VR and neuromodulation therapies for disorders of the bodily self.

Abstract-Although great advances have been made in the design and control of lower extremity prostheses, walking on different terrains, such as ramps or stairs, and transitioning between these terrains remains a major challenge for the field. In order to generalize biomimetic behaviour of active lowerlimb prostheses top-down volitional control is required but has until recently been deemed unfeasible due to the difficulties involved in acquiring an adequate electromyographic (EMG) signal. In this study, we hypothesize that a transtibial amputee can extend the functionality of a hybrid controller, designed for level ground walking, to stair ascent and descent by volitionally modulating powered plantar-flexion of the prosthesis. We here present data illustrating that the participant is able to reproduce ankle push-off behaviour of the intrinsic controller during stair ascent as well as prevent inadvertent push-off during stair descent. Our findings suggest that EMG signal from the residual limb muscles can be used to transition between level-ground walking and stair ascent/descent within a single step and significantly improve prosthesis performance during stair-ambulation.

Sensory attenuation is typically observed for self-generated compared to externally generated action effects. In the present study we investigated whether auditory sensory suppression is modulated as a function of sounds being generated by the upper or lower limbs. We report sensory attenuation, as reflected in a reduced auditory N1 component, which was comparable for sounds generated by the lower and the upper limbs. Increasing temporal delays between actions and sounds did not modulate suppression of the N1 component, but did have an effect on the latency of the N1 component. In contrast, for the P2 component sensory suppression was only observed for sounds generated by the hands and presented at short latencies. These findings provide new insight into the functional and neural dynamics of sensory suppression and suggest the existence of comparable agency mechanisms for both the upper and the lower limbs.

The primary purpose of this study was to investigate the effects of cognitive loading on movement kinematics and trajectory formation during goal-directed walking in a virtual reality (VR) environment. The secondary objective was to measure how participants corrected their trajectories for perturbed feedback and how participants' awareness of such perturbations changed under cognitive loading. We asked 14 healthy young adults to walk towards four different target locations in a VR environment while their movements were tracked and played back in real-time on a large projection screen. In 75% of all trials we introduced angular deviations of ±5° to ±30° between the veridical walking trajectory and the visual feedback. Participants performed a second experimental block under cognitive load (serial-7 subtraction, counter-balanced across participants). We measured walking kinematics (joint-angles, velocity profiles) and motor performance (end-point-compensation, trajectory-deviations). Motor awareness was determined by asking participants to rate the veracity of the feedback after every trial. In-line with previous findings in natural settings, participants displayed stereotypical walking trajectories in a VR environment. Our results extend these findings as they demonstrate that taxing cognitive resources did not affect trajectory formation and deviations although it interfered with the participants' movement kinematics, in particular walking velocity. Additionally, we report that motor awareness was selectively impaired by the secondary task in trials with high perceptual uncertainty. Compared with data on eye and arm movements our findings lend support to the hypothesis that the central nervous system (CNS) uses common mechanisms to govern goal-directed movements, including locomotion. We discuss our results with respect to the use of VR methods in gait control and rehabilitation.