Hand-related Actions Research Articles

Reducing Grip Uncertainty During Initial Prosthetic Hand Use Improves Eye-Hand Coordination and Lowers Mental Workload

The reliance on vision to control a myoelectric prosthesis is cognitively burdensome and contributes to device abandonment. The feeling of uncertainty when gripping an object is thought to be the cause of this overreliance on vision in hand-related actions. We explored if experimentally reducing grip uncertainty alters the visuomotor control and mental workload experienced during initial prosthesis use. In a repeated measures design, twenty-one able-bodied participants took part in a pouring task across three conditions: (a) using their anatomical hand, (b) using a myoelectric prosthetic hand simulator, and (c) using a myoelectric prosthetic hand simulator with Velcro attached to reduce grip uncertainty. Performance, gaze behaviour (using mobile eye-tracking) and self-reported mental workload, was measured. Results showed that using a prosthesis (with or without Velcro) slowed task performance, impaired typical eye-hand coordination and increased mental workload compared to anatomic hand control. However, when using the prosthesis with Velcro, participants displayed better prosthesis control, more effective eye-hand coordination and reduced mental workload compared to when using the prosthesis without Velcro. These positive results indicate that reducing grip uncertainty could be a useful tool for encouraging more effective prosthesis control strategies in the early stages of prosthetic hand learning.

Foot-related/walking macro-affordances are implicitly activated and preferentially guided by the framing distance of the environmental layout

As classically captured in the notion of affordance, the natural environment presents animals with multiple opportunities for action and locomotion appears as the privileged form of action to cover distance in the extrapersonal space/environment. We have recently described a facilitation effect, known as “macro-affordance”, for the execution of walking-related actions in response to distant vs. near objects/locations in the extrapersonal space. However, since the manipulation of distance was coextensive to landmark-objects contained in the environment and to the environmental layout per se, the relative contribution of these two factors remains undetermined. In addition, since the effect was originally described in the context of an incidental priming paradigm, it is still unknown whether it was specifically associated with an implicit coding of environmental distance. Here, across three experiments, we examined the degree to which the “macro-affordance” effect reflects (i) the encoding of environmental vs. landmark-objects’ distance, (ii) the involvement of an implicit vs. controlled system, (iii) a foot-effector specificity. The results showed that the “macro-affordance” effect is more efficiently triggered by the framing distance of the environmental layout (far/wide/panoramic vs. near/close/restricted) rather than of isolated landmark-objects in the environment and that it only emerges when the distance dimension is implicitly processed within the incidental priming paradigm. The results additionally suggested a specificity of the effect for foot- vs. hand-related actions. The present findings suggest that macro-affordances reflect an implicit coding of spatial features of the environmental layout and viewer–environment relationships that preferentially guide a walking-related exploration of the spatial environment.

Automatic coding of environmental distance for walking-related locomotion in the foot-related sensory-motor system: A TMS study on macro-affordances

We have recently described a facilitation effect for the execution of a walking-related action in response to distant objects/locations in the extrapersonal space. Based on the parallelism with the well-known effect of “micro-affordance”, observed during the execution of functionally appropriate hand-related actions towards manipulable objects, we have referred to this effect in terms of “macro-affordance”. Here we used transcranical magnetic stimulation (TMS) to investigate whether a foot-related region located in the human dorsal precuneate cortex plays a causal role in the generation and maintenance of such behavioral effect. This question was addressed by comparing the magnitude of the facilitation effect during an incidental go/no-go task, i.e. advantage for walking-related actions to pictures framing an environment from a far vs. near distance, during three different TMS conditions. The three TMS conditions were collected in all subjects in a randomized order and included stimulation of: i. a foot-related region in the anterior precuneus, ii. a control region in the middle intraparietal sulcus (mIPS), and iii. a sham condition.Enrollment in the TMS protocol was based on analysis of individual performance during a preliminary session conducted using a sham stimulation.TMS was administered at a low frequency range before the beginning of each condition. The results showed that stimulation of the foot-related region in the anterior precuneus produced a significant reduction of the walking-related facilitation effect as compared to both stimulation of the active-control region and the non-active sham stimulation. These findings suggest that the foot-related sensory-motor system directly participates in the process of extraction of the spatial features (i.e. distance) from an environmental scene that are useful for locomotion. More in general, these findings support an automatic coding of environmental affordance or “macro-affordances” in the walking-related sensory-motor system.

Walking-related locomotion is facilitated by the perception of distant targets in the extrapersonal space

The Gibsonian notion of affordance has been massively employed in cognitive sciences to characterize the tight interdependence between hand-related actions, manipulable objects and peripersonal space. A behavioural facilitation effect, indeed, is observed for grasping actions directed to objects located in the ‘reachable’ peripersonal space. Relevantly, this relationship is supported by dedicated neural systems in the brain. The original notion of affordance, however, was directly inspired by real-time interactions between animals and their extended natural environment. Consistently, also the extrapersonal space representation can be significantly modulated by action-related factors, and the brain contains dedicated systems for the representation of topographical space and navigation. Here we examined whether a facilitation effect could be also described for a walking-related action in the far extrapersonal space. To this aim, we employed a go/no-go paradigm requiring subjects to execute a footstep ahead in response to pictures of a virtual reality environment containing objects located at different distances (near, far) and eccentricities (central, peripheral). A walking-related, facilitation effect for distant extrapersonal locations was found, suggesting an automatic trigger of walking by positions that preferentially guide spatial exploration. Based on the parallelism with the literature on micro-affordances, we propose that this effect can be described in terms of “macro-affordances”.

A Double Dissociation in Sensitivity to Verb and Noun Semantics Across Cortical Networks.

What is the neural organization of the mental lexicon? Previous research suggests that partially distinct cortical networks are active during verb and noun processing, but what information do these networks represent? We used multivoxel pattern analysis (MVPA) to investigate whether these networks are sensitive to lexicosemantic distinctions among verbs and among nouns and, if so, whether they are more sensitive to distinctions among words in their preferred grammatical class. Participants heard 4 types of verbs (light emission, sound emission, hand-related actions, mouth-related actions) and 4 types of nouns (birds, mammals, manmade places, natural places). As previously shown, the left posterior middle temporal gyrus (LMTG+), and inferior frontal gyrus (LIFG) responded more to verbs, whereas the inferior parietal lobule (LIP), precuneus (LPC), and inferior temporal (LIT) cortex responded more to nouns. MVPA revealed a double-dissociation in lexicosemantic sensitivity: classification was more accurate among verbs than nouns in the LMTG+, and among nouns than verbs in the LIP, LPC, and LIT. However, classification was similar for verbs and nouns in the LIFG, and above chance for the nonpreferred category in all regions. These results suggest that the lexicosemantic information about verbs and nouns is represented in partially nonoverlapping networks.

Neural Correlates of Semantic Prediction and Resolution in Sentence Processing.

Most brain-imaging studies of language comprehension focus on activity following meaningful stimuli. Testing adult human participants with high-density EEG, we show that, already before the presentation of a critical word, context-induced semantic predictions are reflected by a neurophysiological index, which we therefore call the semantic readiness potential (SRP). The SRP precedes critical words if a previous sentence context constrains the upcoming semantic content (high-constraint contexts), but not in unpredictable (low-constraint) contexts. Specific semantic predictions were indexed by SRP sources within the motor system—in dorsolateral hand motor areas for expected hand-related words (e.g., “write”), but in ventral motor cortex for face-related words (“talk”). Compared with affirmative sentences, negated ones led to medial prefrontal and more widespread motor source activation, the latter being consistent with predictive semantic computation of alternatives to the negated expected concept. Predictive processing of semantic alternatives in negated sentences is further supported by a negative-going event-related potential at ∼400 ms (N400), which showed the typical enhancement to semantically incongruent sentence endings only in high-constraint affirmative contexts, but not to high-constraint negated ones. These brain dynamics reveal the interplay between semantic prediction and resolution (match vs error) processing in sentence understanding.SIGNIFICANCE STATEMENT Most neuroscientists agree on the eminent importance of predictive mechanisms for understanding basic as well as higher brain functions. This contrasts with a sparseness of brain measures that directly reflects specific aspects of prediction, as they are relevant in the processing of language and thought. Here we show that when critical words are strongly expected in their sentence context, a predictive brain response reflects meaning features of these anticipated symbols already before they appear. The granularity of the semantic predictions was so fine grained that the cortical sources in sensorimotor and medial prefrontal cortex even distinguished between predicted face- or hand-related action words (e.g., the words “lick” or “pick”) and between affirmative and negated sentence meanings.

How the context matters. Literal and figurative meaning in the embodied language paradigm.

The involvement of the sensorimotor system in language understanding has been widely demonstrated. However, the role of context in these studies has only recently started to be addressed. Though words are bearers of a semantic potential, meaning is the product of a pragmatic process. It needs to be situated in a context to be disambiguated. The aim of this study was to test the hypothesis that embodied simulation occurring during linguistic processing is contextually modulated to the extent that the same sentence, depending on the context of utterance, leads to the activation of different effector-specific brain motor areas. In order to test this hypothesis, we asked subjects to give a motor response with the hand or the foot to the presentation of ambiguous idioms containing action-related words when these are preceded by context sentences. The results directly support our hypothesis only in relation to the comprehension of hand-related action sentences.

Neuronal bases of peripersonal and extrapersonal spaces, their plasticity and their dynamics: Knowns and unknowns

While space is perceived as unitary, experimental evidence indicates that the brain actually contains a modular representation of space, specific cortical regions being involved in the processing of extra-personal space, that is the space that is far away from the subject and that cannot be directly acted upon by the body, while other cortical regions process peripersonal space, that is the space that directly surrounds us and which we can act upon. In the present review, we focus on non-human primate research and we review the single cells, areal and cortical functional network mechanisms that are proposed to underlie extrapersonal and peripersonal space representations. Importantly, the current dominant framework for the study of peripersonal space is centered on the key notion that actions and specifically arm and hand-related actions, shape cortical peripersonal space representations. In the present review, we propose to enlarge this framework to include other variables that have the potential to shape peripersonal space representations, namely emotional and social information. In the initial section of the manuscript, we thus first provide an extensive up-to-date review of the low level sensory and oculomotor signals that contribute to the construction of a core cortical far and near space representation, in key parietal, premotor and prefrontal periarcuate cortical regions. We then highlight the key functional properties that are needed to encode peripersonal space and we narrow down our discussion to the specific parietal and periarcuate areas that share these properties: the parieto-premotor peripersonal space network and the parieto-premotor network for grasping. Last, we review evidence for a changing peripersonal space representation. While plastic changes in peripersonal space representation have been described during tool use and their underlying neural bases have been well characterized, the description of dynamical changes in peripersonal space representation as a function of the emotional or social context is quite novel and relies on behavioral human studies. The neural bases of such a dynamic adjustments of peripersonal space coding are yet unknown. We thus review these novel observations and we discuss their putative underlying neural bases.

Neuronal correlates of decisions to speak and act: Spontaneous emergence and dynamic topographies in a computational model of frontal and temporal areas

The neural mechanisms underlying the spontaneous, stimulus-independent emergence of intentions and decisions to act are poorly understood. Using a neurobiologically realistic model of frontal and temporal areas of the brain, we simulated the learning of perception–action circuits for speech and hand-related actions and subsequently observed their spontaneous behaviour. Noise-driven accumulation of reverberant activity in these circuits leads to their spontaneous ignition and partial-to-full activation, which we interpret, respectively, as model correlates of action intention emergence and action decision-and-execution. Importantly, activity emerged first in higher-association prefrontal and temporal cortices, subsequently spreading to secondary and finally primary sensorimotor model-areas, hence reproducing the dynamics of cortical correlates of voluntary action revealed by readiness-potential and verb-generation experiments. This model for the first time explains the cortical origins and topography of endogenous action decisions, and the natural emergence of functional specialisation in the cortex, as mechanistic consequences of neurobiological principles, anatomical structure and sensorimotor experience.

The effects of rTMS over the primary motor cortex: The link between action and language

Is the primary motor cortex (M1) necessary for language comprehension? The present study investigates the role of the primary motor cortex during verbs comprehension, within the framework of the embodied theories of language. We applied rTMS over the right and left hand portion of M1 and tested the effects of the stimulation toward the processing of hand-related action verbs versus abstract verbs. Results underlined a specific inhibition effect following left stimulation, only with hand-related action verbs. These findings seem to corroborate the hypothesis of a functional role of M1 in action verbs comprehension.

Hands on the future: facilitation of cortico‐spinal hand‐representation when reading the future tense of hand‐related action verbs

Reading action-related verbs brings about sensorimotor neural activity, suggesting that the linguistic representation of actions impinges upon neural structures largely overlapping with those involved in actual action execution. While studies of direct action observation indicate that motor mirroring is inherently anticipatory, no information is currently available on whether deriving action-related knowledge from language also takes into account the temporal deployment of actions. Using transcranial magnetic stimulation, here we sought to determine whether reading action verbs conjugated in the future induced higher cortico-spinal activity with respect to when the same verbs were conjugated in the past tense. We recorded motor-evoked potentials (MEPs) from relaxed hand and leg muscles of healthy subjects who were reading silently hand- or leg-related action, sensorial (non-somatic) and abstract verbs conjugated either in future or past tense. The amplitude of MEPs recorded from the hand was higher during reading hand-related action verbs conjugated in the future than in the past. No future-related modulation of leg muscles activity was found during reading leg-related action verbs. In a similar vein, no future-related change of hand or leg muscles reactivity was found for abstract or sensorial verbs. These results indicate that the anticipatory mirroring of hand actions may be triggered by linguistic representations and not only by direct action observation.

Action verbs and the primary motor cortex: A comparative TMS study of silent reading, frequency judgments, and motor imagery

Single pulse transcranial magnetic stimulation (TMS) was applied to the hand area of the left primary motor cortex or, as a control, to the vertex (STIMULATION: TMSM1 vs. TMSvertex) while right-handed volunteers silently read verbs related to hand actions. We examined three different tasks and time points for stimulation within the same experiment: subjects indicated with their left foot when they (i) had finished reading, (ii) had judged whether the corresponding movement involved a hand rotation after simulating the hand movement, and (iii) had judged whether they would frequently encounter the action verb in a newspaper (TASK: silent reading, motor imagery, and frequency judgment). Response times were compared between TMSM1 and TMSvertex, both applied at different time points after stimulus onset (DELAY: 150, 300, 450, 600, and 750ms).TMSM1 differentially modulated task performance: there was a significant facilitatory effect of TMSM1 for the imagery task only (about 88ms), with subjects responding about 10% faster (compared to TMSvertex). In contrast, response times for silent reading and frequency judgments were unaffected by TMSM1. No differential effect of the time point of TMSM1 was observed.The differential effect of TMSM1 when subjects performed a motor imagery task (relative to performing silent reading or frequency judgments with the same set of verbs) suggests that the primary motor cortex is critically involved in processing action verbs only when subjects are simulating the corresponding movement. This task-dependent effect of hand motor cortex TMS on the processing of hand-related action verbs is discussed with respect to the notion of embodied cognition and the associationist theory.