Biases in hand perception are driven by somatosensory computations, not a distorted hand model.

To sense and interact with objects in the environment, we effortlessly configure our fingertips at desired locations. It is therefore reasonable to assume that the underlying control mechanisms rely on accurate knowledge about the structure and spatial dimensions of our hand and fingers. This intuition, however, is challenged by years of research showing drastic biases in the perception of finger geometry.1,2,3,4,5 This perceptual bias has been taken as evidence that the brain's internal representation of the body's geometry is distorted,6 leading to an apparent paradox regarding the skillfulness of our actions.7 Here, we propose an alternative explanation of the biases in hand perception-they are the result of the Bayesian integration of noisy, but unbiased, somatosensory signals about finger geometry and posture. To address this hypothesis, we combined Bayesian reverse engineering with behavioral experimentation on joint and fingertip localization of the index finger. We modeled the Bayesian integration either in sensory or in space-based coordinates, showing that the latter model variant led to biases in finger perception despite accurate representation of finger length. Behavioral measures of joint and fingertip localization responses showed similar biases, which were well fitted by the space-based, but not the sensory-based, model variant. The space-based model variant also outperformed a distorted hand model with built-in geometric biases. In total, our results suggest that perceptual distortions of finger geometry do not reflect a distorted hand model but originate from near-optimal Bayesian inference on somatosensory signals.

Peripersonal space around the upper and the lower limbs.

Peripersonal space (PPS), the space closely surrounding the body, is typically characterised by enhanced multisensory integration. Neurophysiological and behavioural studies have consistently shown stronger visuo-tactile integration when a visual stimulus is presented close to the tactually stimulate body part in near space (within PPS) than in far space. However, in the majority of these studies, tactile stimuli were delivered to the upper limbs, torso and face. Therefore, it is not known whether the space surrounding the lower limbs is characterised by similar multisensory properties. To address this question, we asked participants to complete two versions of the classic visuo-tactile crossmodal congruency task in which they had to perform speeded elevation judgements of tactile stimuli presented to the dorsum of the hand and foot while a simultaneous visual distractor was presented at spatially congruent or incongruent locations either in near or far space. In line with existing evidence, when the tactile target was presented to the hand, the size of the crossmodal congruency effect (CCE) decreased in far as compared to near space, suggesting stronger visuo-tactile multisensory integration within PPS. In contrast, when the tactile target was presented to the foot, the CCE decreased for visual distractors in near than far space. These findings show systematic differences between the representation of PPS around upper and lower limbs, suggesting that the multisensory properties of the different body part-centred representations of PPS are likely to depend on the potential actions performed by the different body parts.

Cortical activation and motor body representations in a patient with subacute sclerosing panencephalitis.

The current neuroimaging study investigated the sensorimotor maps during hand, feet and lips movements at one year after diagnosis of of subacute sclerosing panencephalitis (SSPE) in a 17 years-old patient. A lesion prediction algorithm showed that the posterior thalamic radiations, the splenium of the corpus callosum, the posterior and superior corona radiate, and the cingolum, showed a high lesion probability. Comparing the fMRI activations of the left and right hemisphere, we found that the representation of the left hand movement was more inferior/anterior and less represented than the representation of the right one; and the representation of the right foot movement was more superior, less represented than the representation of the left one and poorly activated at the predefined statistical threshold. The fMRI results are in line with the clinical report, describing an asymmetrical distribution of the periodic stereotyped myoclonic jerks, which mainly occurred for the left arm/hand and for the right leg/foot. This is the first fMRI study investigating the representation of the body parts in patients with SSPE. Results show that in SSPE the hyper-stimulation of the motor system (dedicated to the arm/hand and leg/foot more involved by the occurrence of the jerks) is accompanied by an under-activation of the corresponding motor representations in coincidence with voluntary movements.

"Mind the thumb": Judging hand laterality is anchored on the thumb position.

People can decide whether the image of a hand represents a left or a right one. The laterality judgment mainly implies mentally imaging own hand movement (motor simulation) if the stimulus represents a palm, or analysing visual cues, as hand asymmetry, if the stimulus reproduces a dorsum. Here, by capitalizing on evidence underscoring the key role of thumb-palm complex in motor dexterity of human hand, we hypothesise that activation of motor or visual processes when judging hand laterality is due to the different relevance of palm-thumb and dorsum-thumb combinations to hand action. To test this thumb-anchored strategy, in a laterality judgment experiment, we concurrently manipulated the thumb position (flexed or extended) with respect to palm and dorsum, and the human likeness of the hand shape (influencing the salience of the thumb with respect to the hand shape). The main results demonstrated that viewing the flexed thumb from palm or dorsum elicited motor simulation, while viewing the extended thumb activated motor simulation when combined with palm but not dorsum. The present data highlight the pivotal role of the thumb in hand laterality judgments, consistent with its key role in human in-hand manipulation.

Intact tactile anisotropy despite altered hand perception in complex regional pain syndrome: rethinking the role of the primary sensory cortex in tactile and perceptual dysfunction.

Complex Regional Pain Syndrome (CRPS) is characterised by pain, autonomic, sensory and motor abnormalities. It is associated with changes in the primary somatosensory cortex (S1 representation), reductions in tactile sensitivity (tested by two-point discrimination), and alterations in perceived hand size or shape (hand perception). The frequent co-occurrence of these three phenomena has led to the assumption that S1 changes underlie tactile sensitivity and perceptual disturbances. However, studies underpinning such a presumed relationship use tactile sensitivity paradigms that involve the processing of both non-spatial and spatial cues. Here, we used a task that evaluates anisotropy (i.e., orientation-dependency; a feature of peripheral and S1 representation) to interrogate spatial processing of tactile input in CRPS and its relation to hand perception. People with upper limb CRPS (n = 14) and controls with (n = 15) or without pain (n = 19) judged tactile distances between stimuli-pairs applied across and along the back of either hand to provide measures of tactile anisotropy. Hand perception was evaluated using a visual scaling task and questionnaires. Data were analysed with generalised estimating equations. Contrary to our hypotheses, tactile anisotropy was bilaterally preserved in CRPS, and the magnitude of anisotropic perception bias was comparable between groups. Hand perception was distorted in CRPS but not related to the magnitude of anisotropy or bias. Our results suggest against impairments in spatial processing of tactile input, and by implication S1 representation, as the cause of distorted hand perception in CRPS. Further work is warranted to elucidate the mechanisms of somatosensory dysfunction and distorted hand perception in CRPS.

Enhancement of perceived body ownership in virtual reality-based teleoperation may backfire in the execution of high-risk tasks.

Guided by previous research on the role of embodiment in virtual environments, this study aimed to investigate the potential effects of using human-like (compared to robotic) virtual hands on work performances in the context of virtual reality (VR)-based teleoperation of high-risk machinery. A 2 × 2 mixed factorial design experiment (N = 74), with the virtual hand representation as a within-subjects factor (robotic vs. human-like virtual hands) and the risk of danger as a between-subjects factor (low vs. high), was conducted to examine the effects of virtual hand representations (i.e., human-likeness) on perceived body ownership (i.e., embodiment), risk perception, intention to work using the teleoperator, and work performance (i.e., the number of successful task completions). In addition, the moderating effects of the risk of danger on the relationship between perceived body ownership and risk perception were explored. Results showed that the enhancement of perceived body ownership in VR-based teleoperation, induced by the use of human-like hands, increased the risk perception and degraded workers' task performances in the execution of high-risk tasks. Further implications of the findings were discussed.

Tactile distance adaptation aftereffects do not transfer to perceptual hand maps.

Recent studies have demonstrated that mental representations of the hand dorsum are distorted even for healthy participants. Perceptual hand maps estimated by pointing to specific landmarks (e.g., knuckles and tips of fingers) is stretched and shrunk along the medio-lateral and the proximo-distal axes, respectively. Similarly, tactile distance perception between two touches is longer along the medio-lateral axis than the proximo-distal axis. The congruency of the two types of distortions suggests that common perceptual and neural representations may be involved in these processes. Prolonged stimulation by two simultaneous touches having a particular distance can bias subsequent perception of tactile distances (e.g., adaptation to a long distance induces shorter stimuli to be perceived even shorter). This tactile distance adaptation aftereffect has been suggested to occur based on the modulations of perceptual and neural responses at low somatosensory processing stages. The current study investigated whether tactile distance adaptation aftereffects affect also the pattern of distortions on the perceptual hand maps. Participants localized locations on the hand dorsum cued by tactile stimulations (Experiment 1) or visually presented landmarks on a hand silhouette (Experiment 2). Each trial was preceded by adaptation to either a small (2 cm) or large (4 cm) tactile distance. We found clear tactile distance aftereffects. However, no changes were observed for the distorted pattern of the perceptual hand maps following adaptation to a tactile distance. Our results showed that internal body representations involved in perceptual distortions may be distinct between tactile distance perception and the perceptual hand maps underlying position sense.

'Not only faces': specialized visual representation of human hands revealed by adaptation.

Classical neurophysiological studies demonstrated that the monkey brain is equipped with neurons selectively representing the visual shape of the primate hand. Neuroimaging in humans provided data suggesting that a similar representation can be found in humans. Here, we investigated the selectivity of hand representation in humans by means of the visual adaptation technique. Results showed that participants' judgement of human-likeness of a visual probe representing a human hand was specifically reduced by a visual adaptation procedure when using a human hand adaptor but not when using an anthropoid robotic hand or a non-primate animal paw adaptor. Instead, human-likeness of the anthropoid robotic hand was affected by both human and robotic adaptors. No effect was found when using a non-primate animal paw as adaptor or probe. These results support the existence of specific neural mechanisms encoding human hand in the human's visual system.

Preservation of hand movement representation in the sensorimotor areas of amputees.

Denervation due to amputation is known to induce cortical reorganization in the sensorimotor cortex. Although there is evidence that reorganization does not lead to a complete loss of the representation of the phantom limb, it is unclear to what extent detailed, finger-specific activation patterns are preserved in motor cortex, an issue that is also relevant for development of brain-computer interface solutions for paralysed people. We applied machine learning to obtain a quantitative measure for the functional organization within the motor and adjacent cortices in amputees, using high resolution functional MRI and attempted hand gestures. Subjects with above-elbow arm amputation (n = 8) and non-amputated controls (n = 9) made several gestures with either their right or left hand. Amputees attempted to make gestures with their amputated hand. Images were acquired using 7 T functional MRI. The sensorimotor cortex was divided into four regions, and activity patterns were classified in individual subjects using a support vector machine. Classification scores were significantly above chance for all subjects and all hands, and were highly similar between amputees and controls in most regions. Decodability of phantom movements from primary motor cortex reached the levels of right hand movements in controls. Attempted movements were successfully decoded from primary sensory cortex in amputees, albeit lower than in controls but well above chance level despite absence of somatosensory feedback. There was no significant correlation between decodability and years since amputation, or age. The ability to decode attempted gestures demonstrates that the detailed hand representation is preserved in motor cortex and adjacent regions after denervation. This encourages targeting sensorimotor activity patterns for development of brain-computer interfaces.

Modulation of hand motor-related area during motor imagery and motor execution before and after middle 2/5 of the MS6 line scalp acupuncture stimulation: An fMRI study.

Scalp acupuncture (SA) combines the concept of cerebral cortex organization with the principles of acupuncture. The SA stimulates sections of the cerebral cortex. We studied the functional modulation of the left hand sensorimotor area induced by SA in order to investigate the specificity of the SA-related functional effects of the middle 2/5 of the MS6 line of the left side, which corresponds to the upper limb motor segment of the primary motor area. To this purpose, we compared the pre- and post-SA functional activation patterns during an implicit motor imagery task (handedness decision in which participants simulated rotational hand movements) and an explicit manual motor execution task. Feet and mouth movements, and the fMRI changes in their respective representations were used as control conditions. Only SA on the hand area of the left side (as compared to the mouth and the foot representations which were used as control conditions) exerted a release effect on the right hand area. In addition, an increased activation of the superior parietal lobe was seen, which is involved in movement control and planning. Taken together, these preliminary findings may shed light on the SA effects and confirm a prolonged effect of SA even after cessation of needling stimulation.

Cortical connections to single digit representations in area 3b of somatosensory cortex in squirrel monkeys and prosimian galagos.

The ventral posterior nucleus of thalamus sends highly segregated inputs into each digit representation in area 3b of primary somatosensory cortex. However, the spatial organization of the connections that link digit representations of areas 3b with other somatosensory areas is less understood. Here we examined the cortical inputs to individual digit representations of area 3b in four squirrel monkeys and one prosimian galago. Retrograde tracers were injected into neurophysiologically defined representations of individual digits of area 3b. Cortical tissues were cut parallel to the surface in some cases and showed that feedback projections to individual digits overlapped extensively in the hand representations of areas 3b, 1, and parietal ventral (PV) and second somatosensory (S2) areas. Other regions with overlapping populations of labeled cells included area 3a and primary motor cortex (M1). The results were confirmed in other cases in which the cortical tissues were cut in the coronal plane. The same cases also showed that cells were primarily labeled in the infragranular and supragranular layers. Thus, feedback projections to individual digit representations in area 3b mainly originate from multiple digits and other portions of hand representations of areas 3b, 1, PV, and S2. This organization is in stark contrast to the segregated thalamocortical inputs, which originate in single digit representations and terminate in the matching digit representation in the cortex. The organization of feedback connections could provide a substrate for the integration of information across the representations of adjacent digits in area 3b.