Helge Gillmeister

Through the looking glass: Counter-mirror activation following incompatible sensorimotor learning

The mirror system, comprising cortical areas that allow the actions of others to be represented in the observer’s own motor system, is thought to be crucial for the development of social cognition in humans. Despite the importance of the human mirror system, little is known about its origins. We investigated the role of sensorimotor experience in the development of the mirror system. Functional magnetic resonance imaging was used to measure neural responses to observed hand and foot actions following one of two types of training. During training, participants in the Compatible (control) group made mirror responses to observed actions (hand responses were made to hand stimuli and foot responses to foot stimuli), whereas the Incompatible group made counter‐mirror responses (hand to foot and foot to hand). Comparison of these groups revealed that, after training to respond in a counter‐mirror fashion, the relative action observation properties of the mirror system were reversed; areas that showed greater responses to observation of hand actions in the Compatible group responded more strongly to observation of foot actions in the Incompatible group. These results suggest that, rather than being innate or the product of unimodal visual or motor experience, the mirror properties of the mirror system are acquired through sensorimotor learning.

Sensorimotor experience enhances automatic imitation of robotic action.

Recent research in cognitive neuroscience has found that observation of human actions activates the ‘mirror system’ and provokes automatic imitation to a greater extent than observation of non-biological movements. The present study investigated whether this human bias depends primarily on phylogenetic or ontogenetic factors by examining the effects of sensorimotor experience on automatic imitation of non-biological robotic, stimuli. Automatic imitation of human and robotic action stimuli was assessed before and after training. During these test sessions, participants were required to execute a pre-specified response (e.g. to open their hand) while observing a human or robotic hand making a compatible (opening) or incompatible (closing) movement. During training, participants executed opening and closing hand actions while observing compatible (group CT) or incompatible movements (group IT) of a robotic hand. Compatible, but not incompatible, training increased automatic imitation of robotic stimuli (speed of responding on compatible trials, compared with incompatible trials) and abolished the human bias observed at pre-test. These findings suggest that the development of the mirror system depends on sensorimotor experience, and that, in our species, it is biased in favour of human action stimuli because these are more abundant than non-biological action stimuli in typical developmental environments.

Experience-based priming of body parts: a study of action imitation.

Two important dimensions of action are the movement and the body part with which the movement is effected. Experiment 1 tested whether automatic imitation is sensitive to the body part dimension of action. We found that hand and foot movements were selectively primed by observation of a corresponding, task-irrelevant effector in motion. Experiment 2 used this body part priming effect to investigate the role of sensorimotor learning in the development of imitation. The results showed that incompatible training, in which observation of hand movements was paired with execution of foot movements and vice versa, led to a greater reduction in body part priming than compatible training, in which subjects experienced typical contingencies between observation and execution of hand and foot movements. These findings are consistent with the assumption that overt behavioral imitation is mediated by the mirror neuron system, which is somatotopically organized. Our results also support the hypothesis that the development of imitation and the mirror neuron system are driven by correlated sensorimotor learning.

Tactile enhancement of auditory detection and perceived loudness.

To study the effects of touch on auditory processing, we examined whether uninformative and irrelevant tactile stimuli presented together with task-relevant sounds can improve auditory detection (Experiment 1), and enhance perceived loudness (Experiment 2). We demonstrated that irrelevant tactile signals facilitate the detection of faint tones, and increase auditory intensity ratings. These crossmodal facilitation effects were found for synchronous when compared to asynchronous auditory-tactile stimulation, and were stronger for weaker than for louder sounds. They are interpreted in terms of a multisensory integration mechanism that increases the strength of auditory signals, and adheres to the rules of inverse effectiveness and temporal (but not spatial) co-occurrence. This integration might be mediated by auditory-tactile multisensory neurons in regions of auditory association cortex that are also involved in auditory detection and loudness discrimination.

Viewing the body modulates neural mechanisms underlying sustained spatial attention in touch

Cross‐modal links between vision and touch have been extensively shown with a variety of paradigms. The present event‐related potential (ERP) study aimed to clarify whether neural mechanisms underlying sustained tactile‐spatial attention may be modulated by visual input, and the sight of the stimulated body part (i.e. hands) in particular. Participants covertly attended to one of their hands throughout a block to detect infrequent tactile target stimuli at that hand while ignoring tactile targets at the unattended hand, and all tactile non‐targets. In different blocks, participants performed this task under three viewing conditions: full vision; hands covered from view; and blindfolded. When the participants’ hands were visible attention was found to modulate somatosensory ERPs at early latencies (i.e. in the time range of the somatosensory P100 and the N140 components), as well as at later time intervals, from 200 ms after stimulus onset. By contrast, when participants were blindfolded and, crucially, even when only their hands were not visible, attentional modulations were found to arise only at later intervals (i.e. from 200 ms post‐stimulus), while earlier somatosensory components were not affected by spatial attention. The behavioural results tallied with these electrophysiological findings, showing faster response times to tactile targets under the full vision condition compared with conditions when participants’ hands were covered, and when participants were blindfolded. The results from this study provide the first evidence of the profound impact of vision on mechanisms underlying sustained tactile‐spatial attention, which is enhanced by the sight of the body parts (i.e. hands).

Which finger? Early effects of attentional selection within the hand are absent when the hand is viewed

The sight of a hand can bias the distribution of spatial attention, and recently it has been shown that viewing both hands simultaneously can facilitate spatial selection between tactile events at the hands when these are far apart. Here we directly compared the electrophysiological correlates of within‐hand and between‐hands tactile–spatial selection to investigate whether within‐hand selection is similarly facilitated by viewing the fingers. Using somatosensory event‐related potentials, we have shown that effects of selection between adjacent fingers of the same hand at early somatosensory components P45 and N80 were absent when the fingers were viewed. Thus, we found a detrimental effect of vision on tactile–spatial within‐body part (i.e. hand) selection. In contrast, effects of tactile–spatial selection between hands placed next to each other, which were first found at the P100 component, were unaffected by vision of the hands. Our findings suggest that (i) within‐hand and between‐hands selection can operate at different stages of processing, and (ii) the effects of vision on within‐hand and between‐hands attentional selection may reflect fundamentally different mechanisms.

Hands behind your back: effects of arm posture on tactile attention in the space behind the body

Previous research has shown that tactile-spatial information originating from the front of the body is remapped from an anatomical to an external spatial coordinate system, guided by the availability of visual information early in development. Comparably little is known about regions of space for which visual information is not typically available, such as the space behind the body. This study tests for the first time the electrophysiological correlates of the effects of proprioceptive information on tactile-attentional mechanisms in the space behind the back. Observers were blindfolded and tactually cued to detect infrequent tactile targets on either their left or right hand and to respond to them either vocally or with index finger movements. We measured event-related potentials to tactile probes on the hands in order to explore tactile-spatial attention when the hands were either held close together or far apart behind the observer’s back. Results show systematic effects of arm posture on tactile-spatial attention different from those previously found for front space. While attentional selection is typically more effective for hands placed far apart than close together in front space, we found that selection occurred more rapidly for close than far hands behind the back, during both covert attention and movement preparation tasks. This suggests that proprioceptive space may “wrap” around the body, following the hands as they extend horizontally from the front body midline to the center of the back.

ERP investigation of transient attentional selection of single and multiple locations within touch

Mechanisms underlying pure tactile attentional selection were investigated. Tactile imperative stimuli were preceded by symbolic tactile cues directing attention to the left or right (directional cues), or to both hands (non-directional cues). Comparison of ERP waveforms on directional and non-directional cue trials showed that attentional modulations at N140 and P200 components reflect mainly enhancement of stimuli at the attended, while longer latency modulations reflect mainly suppression of processing of stimuli at the unattended location. This pattern of results differs from analogous studies involving other modalities suggesting that different mechanisms underlie pure tactile attention. Furthermore, ERP waveforms on non-directional cue trials were enhanced in comparison to directional cue trials at the P100 component and at longer latencies, indicating that tactile attentional mechanisms may differ when attending to one compared to multiple locations.

Inter-Individual Differences in Vicarious Tactile Perception: a View Across the Lifespan in Typical and Atypical Populations

Touch is our most interpersonal sense, and so it stands to reason that we represent not only our own bodily experiences, but also those felt by others. This review will summarise brain and behavioural research on vicarious tactile perception (mirror touch). Specifically, we will focus on vicarious touch across the lifespan in typical and atypical groups, and will identify the knowledge gaps that are in urgent need of filling by examining what is known about how individuals differ within and between typical and atypical groups.

A new perceptual paradigm to investigate the visual remapping of others’ tactile sensations onto one’s own body shows “mirror touch” for the hands

The last two decades have seen a multitude of publications showing the activation of an observer’s somatosensory cortical system during the observation of touch on another person. Behavioral demonstrations of “mirror touch,” however, have been slow in coming forward, and have so far primarily been shown as “visual remapping of touch” on the face. The present study uses a new paradigm to investigate the mirroring of others’ tactile sensations: a 2-AFC task of intensity judgment for touch on the observer’s left and right index finger pads. Observers viewed a left and right hand in an egocentric position, which were either touched passively (pencil moving to touch index finger pad) or actively sought touch (index finger moving to touch pencil). Touch and no-touch events for the two viewed hands were designed to eliminate confounding effects of spatial attention. Felt touches were either concurrent with viewed touch or no-touch events, or were delayed in time to assess potential response bias. The findings demonstrate visual remapping of touch for touch on the hands. If touch was shown on one of the hands only (e.g., left), observers were more likely to perceive touch on the same hand (i.e., their own left hand) as more intense than touch on the other hand even if tactile intensities did not differ, compared to touch shown on both or neither hand. These remapping effects occurred only when viewed and felt touches were concurrent, they were strongly modulated by the way in which viewed touch was incurred, and they were more reliable for touch on the left hand. A second, control experiment, in which touch observation was replaced by bright dots shown on or next to the finger pads, confirmed that these effects were largely due to genuine tactile mirroring rather than to somatotopic cueing. This 2-AFC tactile intensity judgment task may be a useful paradigm to investigate the remapping of others’ tactile sensations onto an observer’s own body.