The Representation of Observed Actions at the Subordinate, Basic, and Superordinate Level.

Actions can be planned and recognized at different hierarchical levels, ranging from very specific (e.g., to swim backstroke) to very broad (e.g., locomotion). Understanding the corresponding neural representation is an important prerequisite to reveal how our brain flexibly assigns meaning to the world around us. To address this question, we conducted an event-related fMRI study in male and female human participants in which we examined distinct representations of observed actions at the subordinate, basic and superordinate level. Using multiple regression representational similarity analysis (RSA) in predefined regions of interest, we found that the three different taxonomic levels were best captured by patterns of activations in bilateral lateral occipitotemporal cortex (LOTC), showing the highest similarity with the basic level model. A whole-brain multiple regression RSA revealed that information unique to the basic level was captured by patterns of activation in dorsal and ventral portions of the LOTC and in parietal regions. By contrast, the unique information for the subordinate level was limited to bilateral occipitotemporal cortex, while no single cluster was obtained that captured unique information for the superordinate level. The behaviorally established action space was best captured by patterns of activation in the LOTC and superior parietal cortex, and the corresponding neural patterns of activation showed the highest similarity with patterns of activation corresponding to the basic level model. Together, our results suggest that occipitotemporal cortex shows a preference for the basic level model, with flexible access across the subordinate and the basic level.SIGNIFICANCE STATEMENT The human brain captures information at varying levels of abstraction. It is debated which brain regions host representations across different hierarchical levels, with some studies emphasizing parietal and premotor regions, while other studies highlight the role of the lateral occipitotemporal cortex (LOTC). To shed light on this debate, here we examined the representation of observed actions at the three taxonomic levels suggested by Rosch et al. (1976) Our results highlight the role of the LOTC, which hosts a shared representation across the subordinate and the basic level, with the highest similarity with the basic level model. These results shed new light on the hierarchical organization of observed actions and provide insights into the neural basis underlying the basic level advantage.

Overlapping representations of observed actions and action-related features.

The lateral occipitotemporal cortex (LOTC) has been shown to capture the representational structure of a smaller range of actions. In the current study, we carried out an fMRI experiment in which we presented human participants with images depicting 100 different actions and used representational similarity analysis (RSA) to determine which brain regions capture the semantic action space established using judgments of action similarity. Moreover, to determine the contribution of a wide range of action-related features to the neural representation of the semantic action space we constructed an action feature model on the basis of ratings of 44 different features. We found that the semantic action space model and the action feature model are best captured by overlapping activation patterns in bilateral LOTC and ventral occipitotemporal cortex (VOTC). An RSA on eight dimensions resulting from principal component analysis carried out on the action feature model revealed partly overlapping representations within bilateral LOTC, VOTC, and the parietal lobe. Our results suggest spatially overlapping representations of the semantic action space of a wide range of actions and the corresponding action-related features. Together, our results add to our understanding of the kind of representations along the LOTC that support action understanding.

The cognitive structure underlying the organization of observed actions.

In daily life, we frequently encounter actions performed by other people. Here we aimed to examine the key categories and features underlying the organization of a wide range of actions in three behavioral experiments (N = 378 participants). In Experiment 1, we used a multi-arrangement task of 100 different actions. Inverse multidimensional scaling and hierarchical clustering revealed 11 action categories, including Locomotion, Communication, and Aggressive actions. In Experiment 2, we used a feature-listing paradigm to obtain a wide range of action features that were subsequently reduced to 59 key features and used in a rating study (Experiment 3). A direct comparison of the feature ratings obtained in Experiment 3 between actions belonging to the categories identified in Experiment 1 revealed a number of features that appear to be critical for the distinction between these categories, e.g., the features Harm and Noise for the category Aggressive actions, and the features Targeting a person and Contact with others for the category Interaction. Finally, we found that a part of the category-based organization is explained by a combination of weighted features, whereas a significant proportion of variability remained unexplained, suggesting that there are additional sources of information that contribute to the categorization of observed actions. The characterization of action categories and their associated features serves as an important extension of previous studies examining the cognitive structure of actions. Moreover, our results may serve as the basis for future behavioral, neuroimaging and computational modeling studies.

Embodied cognition and mirror neurons: a critical assessment.

According to embodied cognition theories, higher cognitive abilities depend on the reenactment of sensory and motor representations. In the first part of this review, we critically analyze the central claims of embodied theories and argue that the existing behavioral and neuroimaging data do not allow investigators to discriminate between embodied cognition and classical cognitive accounts, which assume that conceptual representations are amodal and symbolic. In the second part, we review the main claims and the core electrophysiological findings typically cited in support of the mirror neuron theory of action understanding, one of the most influential examples of embodied cognition theories. In the final part, we analyze the claim that mirror neurons subserve action understanding by mapping visual representations of observed actions on motor representations, trying to clarify in what sense the representations carried by these neurons can be claimed motor.

The characterization of actions at the superordinate, basic and subordinate level.

Objects can be categorized at different levels of abstraction, ranging from the superordinate (e.g., fruit) and the basic (e.g., apple) to the subordinate level (e.g., golden delicious). The basic level is assumed to play a key role in categorization, e.g., in terms of the number of features used to describe these actions and the speed of processing. To which degree do these principles also apply to the categorization of observed actions? To address this question, we first selected a range of actions at the superordinate (e.g., locomotion), basic (e.g., to swim) and subordinate level (e.g., to swim breaststroke), using verbal material (Experiments 1-3). Experiments 4-6 aimed to determine the characteristics of these actions across the three taxonomic levels. Using a feature listing paradigm (Experiment 4), we determined the number of features that were provided by at least six out of twenty participants (common features), separately for the three different levels. In addition, we examined the number of shared (i.e., provided for more than one category) and distinct (i.e., provided for one category only) features. Participants produced the highest number of common features for actions at the basic level. Actions at the subordinate level shared more features with other actions at the same level than those at the superordinate level. Actions at the superordinate and basic level were described with more distinct features compared to those provided at the subordinate level. Using an auditory priming paradigm (Experiment 5), we observed that participants responded faster to action images preceded by a matching auditory cue corresponding to the basic and subordinate level, but not for superordinate level cues, suggesting that the basic level is the most abstract level at which verbal cues facilitate the processing of an upcoming action. Using a category verification task (Experiment 6), we found that participants were faster and more accurate to verify action categories (depicted as images) at the basic and subordinate level in comparison to the superordinate level. Together, in line with the object categorization literature, our results suggest that information about action categories is maximized at the basic level.

Decoding category and familiarity information during visual imagery.

Visual imagery relies on a widespread network of brain regions, partly engaged during the perception of external stimuli. Beyond the recruitment of category-selective areas (FFA, PPA), perception of familiar faces and places has been reported to engage brain areas associated with semantic information, comprising the precuneus, temporo-parietal junction (TPJ), medial prefrontal cortex (mPFC) and posterior cingulate cortex (PCC). Here we used multivariate pattern analyzes (MVPA) to examine to which degree areas of the visual imagery network, category-selective and semantic areas contain information regarding the category and familiarity of imagined stimuli. Participants were instructed via auditory cues to imagine personally familiar and unfamiliar stimuli (i.e. faces and places). Using region-of-interest (ROI)-based MVPA, we were able to distinguish between imagined faces and places within nodes of the visual imagery network (V1, SPL, aIPS), within category-selective inferotemporal regions (FFA, PPA) and across all brain regions of the extended semantic network (i.e. precuneus, mPFC, IFG and TPJ). Moreover, we were able to decode familiarity of imagined stimuli in the SPL and aIPS, and in some regions of the extended semantic network (in particular, right precuneus, right TPJ), but not in V1. Our results suggest that posterior visual areas - including V1 - host categorical representations about imagined stimuli, and that stimulus familiarity might be an additional aspect that is shared between perception and visual imagery.

Hierarchical Action Encoding Within the Human Brain.

Humans are able to interact with objects with extreme flexibility. To achieve this ability, the brain does not only control specific muscular patterns, but it also needs to represent the abstract goal of an action, irrespective of its implementation. It is debated, however, how abstract action goals are implemented in the brain. To address this question, we used multivariate pattern analysis of functional magnetic resonance imaging data. Human participants performed grasping actions (precision grip, whole hand grip) with two different wrist orientations (canonical, rotated), using either the left or right hand. This design permitted to investigate a hierarchical organization consisting of three levels of abstraction: 1) "concrete action" encoding; 2) "effector-dependent goal" encoding (invariant to wrist orientation); and 3) "effector-independent goal" encoding (invariant to effector and wrist orientation). We found that motor cortices hosted joint encoding of concrete actions and of effector-dependent goals, while the parietal lobe housed a convergence of all three representations, comprising action goals within and across effectors. The left lateral occipito-temporal cortex showed effector-independent goal encoding, but no convergence across the three levels of representation. Our results support a hierarchical organization of action encoding, shedding light on the neural substrates supporting the extraordinary flexibility of human hand behavior.

The Large-Scale Organization of Gestures and Words in the Middle Temporal Gyrus.

The middle temporal gyrus (MTG) has been shown to be recruited during the processing of words, but also during the observation of actions. Here we investigated how information related to words and gestures is organized along the MTG. To this aim, we measured the BOLD response in the MTG to video clips of gestures and spoken words in 17 healthy human adults (male and female). Gestures consisted of videos of an actress performing object-use pantomimes (iconic representations of object-directed actions; e.g., playing guitar), emblems (conventional gestures, e.g., thumb up), and meaningless gestures. Word stimuli (verbs, nouns) consisted of video clips of the same actress pronouncing words. We found a stronger response to meaningful compared with meaningless gestures along the whole left and large portions of the right MTG. Importantly, we observed a gradient, with posterior regions responding more strongly to gestures (pantomimes and emblems) than words and anterior regions showing a stronger response to words than gestures. In an intermediate region in the left hemisphere, the response was significantly higher to words and emblems (i.e., items with a greater arbitrariness of the sign-to-meaning mapping) than to pantomimes. These results show that the large-scale organization of information in the MTG is driven by the input modality and may also reflect the arbitrariness of the relationship between sign and meaning.SIGNIFICANCE STATEMENT Here we investigated the organizing principle of information in the middle temporal gyrus, taking into consideration the input-modality and the arbitrariness of the relationship between a sign and its meaning. We compared the middle temporal gyrus response during the processing of pantomimes, emblems, and spoken words. We found that posterior regions responded more strongly to pantomimes and emblems than to words, whereas anterior regions responded more strongly to words than to pantomimes and emblems. In an intermediate region, only in the left hemisphere, words and emblems evoked a stronger response than pantomimes. Our results identify two organizing principles of neural representation: the modality of communication (gestural or verbal) and the (arbitrariness of the) relationship between sign and meanings.

Visual imagery during real-time fMRI neurofeedback from occipital and superior parietal cortex.

Visual imagery has been suggested to recruit occipital cortex via feedback projections from fronto-parietal regions, suggesting that these feedback projections might be exploited to boost recruitment of occipital cortex by means of real-time neurofeedback. To test this prediction, we instructed a group of healthy participants to perform peripheral visual imagery while they received real-time auditory feedback based on the BOLD signal from either early visual cortex or the medial superior parietal lobe. We examined the amplitude and temporal aspects of the BOLD response in the two regions. Moreover, we compared the impact of self-rated mental focus and vividness of visual imagery on the BOLD responses in these two areas. We found that both early visual cortex and the medial superior parietal cortex are susceptible to auditory neurofeedback within a single feedback session per region. However, the signal in parietal cortex was sustained for a longer time compared to the signal in occipital cortex. Moreover, the BOLD signal in the medial superior parietal lobe was more affected by focus and vividness of the visual imagery than early visual cortex. Our results thus demonstrate that (a) participants can learn to self-regulate the BOLD signal in early visual and parietal cortex within a single session, (b) that different nodes in the visual imagery network respond differently to neurofeedback, and that (c) responses in parietal, but not in occipital cortex are susceptible to self-rated vividness of mental imagery. Together, these results suggest that medial superior parietal cortex might be a suitable candidate to provide real-time feedback to patients suffering from visual field defects.

Directional tuning for eye and arm movements in overlapping regions in human posterior parietal cortex.

A network of frontal and parietal regions is known to be recruited during the planning and execution of arm and eye movements. While movements of the two effectors are typically coupled with each other, it remains unresolved how information is shared between them. Here we aimed to identify regions containing neuronal populations that show directional tuning for both arm and eye movements. In two separate fMRI experiments, the same participants were scanned while performing a center-out arm or eye movement task. Using a whole-brain searchlight-based representational similarity analysis (RSA), we found that a bilateral region in the posterior superior parietal lobule represents both arm and eye movement direction, thus extending previous findings in monkeys.

A norming study of high-quality video clips of pantomimes, emblems, and meaningless gestures.

Recent years have witnessed a growing interest in behavioral and neuroimaging studies on the processing of symbolic communicative gestures, such as pantomimes and emblems, but well-controlled stimuli have been scarce. This study describes a dataset of more than 200 video clips of an actress performing pantomimes (gestures that mimic object-directed/object-use actions; e.g., playing guitar), emblems (conventional gestures; e.g., thumbs up), and meaningless gestures. Gestures were divided into four lists. For each of these four lists, 50 Italian and 50 American raters judged the meaningfulness of the gestures and provided names and descriptions for them. The results of these rating and norming measures are reported separately for the Italian and American raters, offering the first normed set of meaningful and meaningless gestures for experimental studies. The stimuli are available for download via the Figshare database.

Action Categories in Lateral Occipitotemporal Cortex Are Organized Along Sociality and Transitivity.

How neural specificity for distinct conceptual knowledge categories arises is central for understanding the organization of semantic memory in the human brain. Although there is a large body of research on the neural processing of distinct object categories, the organization of action categories remains largely unknown. In particular, it is unknown whether different action categories follow a specific topographical organization on the cortical surface analogously to the category-specific organization of object knowledge. Here, we tested whether the neural representation of action knowledge is organized in terms of nonsocial versus social and object-unrelated versus object-related actions (sociality and transitivity, respectively, hereafter). We hypothesized a major distinction of sociality and transitivity along dorsal and ventral lateral occipitotemporal cortex (LOTC), respectively. Using fMRI-based multivoxel pattern analysis, we identified neural representations of action information associated with sociality and transitivity in bilateral LOTC. Representational similarity analysis revealed a dissociation between dorsal and ventral LOTC. We found that action representations in dorsal LOTC are segregated along features of sociality, whereas action representations in ventral LOTC are segregated along features of transitivity. In addition, representations of sociality and transitivity features were found more anteriorly in LOTC than representations of specific subtypes of actions, suggesting a posterior-anterior gradient from concrete to abstract action features. These findings elucidate how the neural representations of perceptually and conceptually diverse actions are organized in distinct subsystems in the LOTC. SIGNIFICANCE STATEMENT: The lateral occipitotemporal cortex (LOTC) is critically involved in the recognition of objects and actions, but our knowledge about the underlying organizing principles is limited. Here, we discovered a dorsal-ventral distinction of actions in LOTC: dorsal LOTC represents actions based on sociality (how much an action is directed to another person) in proximity to person knowledge. In contrast, ventral LOTC represents actions based on transitivity (how much an action involves the interaction with inanimate objects) in proximity to tools/artifacts in ventral LOTC, suggesting a mutually dependent organization of actions and objects. In addition, we found a posterior-to-anterior organization of the LOTC for concrete and abstract representations, respectively. Our findings provide important insights about the organization of actions in LOTC.

Spatiotopic updating across saccades revealed by spatially-specific fMRI adaptation.

Brain representations of visual space are predominantly eye-centred (retinotopic) yet our experience of the world is largely world-centred (spatiotopic). A long-standing question is how the brain creates continuity between these reference frames across successive eye movements (saccades). Here we use functional magnetic resonance imaging (fMRI) to address whether spatially specific repetition suppression (RS) is evident during trans-saccadic perception. We presented two successive Gabor patches (S1 and S2) in either the upper or lower visual field, left or right of fixation. Spatial congruency was manipulated by having S1 and S2 occur in the same or different upper/lower visual field. On half the trials, a saccade was cued between S1 and S2, placing spatiotopic and retinotopic reference frames in opposition. Equivalent RS was observed in the posterior parietal cortex and frontal eye fields when S1-S2 were spatiotopically congruent, irrespective of whether retinotopic and spatiotopic coordinates were in accord or were placed in opposition by a saccade. Additionally the post-saccadic response to S2 demonstrated spatially-specific RS in retinotopic visual regions, with stronger RS in extrastriate than striate cortex. Collectively, these results are consistent with a robust trans-saccadic spatial updating mechanism for object position that directly influences even the earliest levels of visual processing.

Decoding Concrete and Abstract Action Representations During Explicit and Implicit Conceptual Processing.

Action understanding requires a many-to-one mapping of perceived input onto abstract representations that generalize across concrete features. It is debated whether such abstract action concepts are encoded in ventral premotor cortex (PMv; motor hypothesis) or, alternatively, are represented in lateral occipitotemporal cortex (LOTC; cognitive hypothesis). We used fMRI-based multivoxel pattern analysis to decode observed actions at concrete and abstract, object-independent levels of representation. Participants observed videos of 2 actions involving 2 different objects, using either an explicit or implicit task with respect to conceptual action processing. We decoded concrete action representations by training and testing a classifier to discriminate between actions within each object category. To identify abstract action representations, we trained the classifier to discriminate actions in one object and tested the classifier on actions performed on the other object, and vice versa. Region-of-interest and searchlight analyses revealed decoding in LOTC at both concrete and abstract levels during both tasks, whereas decoding in PMv was restricted to the concrete level during the explicit task. In right inferior parietal cortex, decoding was significant for the abstract level during the explicit task. Our findings are incompatible with the motor hypothesis, but support the cognitive hypothesis of action understanding.

Decoding Internally and Externally Driven Movement Plans.

During movement planning, brain activity within parietofrontal networks encodes information about upcoming actions that can be driven either externally (e.g., by a sensory cue) or internally (i.e., by a choice/decision). Here we used multivariate pattern analysis (MVPA) of fMRI data to distinguish between areas that represent (1) abstract movement plans that generalize across the way in which these were driven, (2) internally driven movement plans, or (3) externally driven movement plans. In a delayed-movement paradigm, human volunteers were asked to plan and execute three types of nonvisually guided right-handed reaching movements toward a central target object: using a precision grip, a power grip, or touching the object without hand preshaping. On separate blocks of trials, movements were either instructed via color cues (Instructed condition), or chosen by the participant (Free-Choice condition). Using ROI-based and whole-brain searchlight-based MVPA, we found abstract representations of planned movements that generalize across the way these movements are selected (internally vs externally driven) in parietal cortex, dorsal premotor cortex, and primary motor cortex contralateral to the acting hand. In addition, we revealed representations specific for internally driven movement plans in contralateral ventral premotor cortex, dorsolateral prefrontal cortex, supramarginal gyrus, and in ipsilateral posterior parietotemporal regions, suggesting that these regions are recruited during movement selection. Finally, we observed representations of externally driven movement plans in bilateral supplementary motor cortex and a similar trend in presupplementary motor cortex, suggesting a role in stimulus-response mapping. SIGNIFICANCE STATEMENT: The way the human brain prepares the body for action constitutes an essential part of our ability to interact with our environment. Previous studies demonstrated that patterns of neuronal activity encode upcoming movements. Here we used multivariate pattern analysis of human fMRI data to distinguish between brain regions containing movement plans for instructed (externally driven) movements, areas involved in movement selection (internally driven), and areas containing abstract movement plans that are invariant to the way these were generated (i.e., that generalize across externally and internally driven movement plans). Our findings extend our understanding of the neural basis of movement planning and have the potential to contribute to the development of brain-controlled neural prosthetic devices.

Decoding actions at different levels of abstraction.

Brain regions that mediate action understanding must contain representations that are action specific and at the same time tolerate a wide range of perceptual variance. Whereas progress has been made in understanding such generalization mechanisms in the object domain, the neural mechanisms to conceptualize actions remain unknown. In particular, there is ongoing dissent between motor-centric and cognitive accounts whether premotor cortex or brain regions in closer relation to perceptual systems, i.e., lateral occipitotemporal cortex, contain neural populations with such mapping properties. To date, it is unclear to which degree action-specific representations in these brain regions generalize from concrete action instantiations to abstract action concepts. However, such information would be crucial to differentiate between motor and cognitive theories. Using ROI-based and searchlight-based fMRI multivoxel pattern decoding, we sought brain regions in human cortex that manage the balancing act between specificity and generality. We investigated a concrete level that distinguishes actions based on perceptual features (e.g., opening vs closing a specific bottle), an intermediate level that generalizes across movement kinematics and specific objects involved in the action (e.g., opening different bottles with cork or screw cap), and an abstract level that additionally generalizes across object category (e.g., opening bottles or boxes). We demonstrate that the inferior parietal and occipitotemporal cortex code actions at abstract levels whereas the premotor cortex codes actions at the concrete level only. Hence, occipitotemporal, but not premotor, regions fulfill the necessary criteria for action understanding. This result is compatible with cognitive theories but strongly undermines motor theories of action understanding.

MEG Multivariate Analysis Reveals Early Abstract Action Representations in the Lateral Occipitotemporal Cortex.

Understanding other people's actions is a fundamental prerequisite for social interactions. Whether action understanding relies on simulating the actions of others in the observers' motor system or on the access to conceptual knowledge stored in nonmotor areas is strongly debated. It has been argued previously that areas that play a crucial role in action understanding should (1) distinguish between different actions, (2) generalize across the ways in which actions are performed (Dinstein et al., 2008; Oosterhof et al., 2013; Caramazza et al., 2014), and (3) have access to action information around the time of action recognition (Hauk et al., 2008). Whereas previous studies focused on the first two criteria, little is known about the dynamics underlying action understanding. We examined which human brain regions are able to distinguish between pointing and grasping, regardless of reach direction (left or right) and effector (left or right hand), using multivariate pattern analysis of magnetoencephalography data. We show that the lateral occipitotemporal cortex (LOTC) has the earliest access to abstract action representations, which coincides with the time point from which there was enough information to allow discriminating between the two actions. By contrast, precentral regions, though recruited early, have access to such abstract representations substantially later. Our results demonstrate that in contrast to the LOTC, the early recruitment of precentral regions does not contain the detailed information that is required to recognize an action. We discuss previous theoretical claims of motor theories and how they are incompatible with our data. SIGNIFICANCE STATEMENT: It is debated whether our ability to understand other people's actions relies on the simulation of actions in the observers' motor system, or is based on access to conceptual knowledge stored in nonmotor areas. Here, using magnetoencephalography in combination with machine learning, we examined where in the brain and at which point in time it is possible to distinguish between pointing and grasping actions regardless of the way in which they are performed (effector, reach direction). We show that, in contrast to the predictions of motor theories of action understanding, the lateral occipitotemporal cortex has access to abstract action representations substantially earlier than precentral regions.

Beta band modulations underlie action representations for movement planning.

To be able to interact with our environment, we need to transform incoming sensory information into goal-directed motor outputs. Whereas our ability to plan an appropriate movement based on sensory information appears effortless and simple, the underlying brain dynamics are still largely unknown. Here we used magnetoencephalography (MEG) to investigate this issue by recording brain activity during the planning of non-visually guided reaching and grasping actions, performed with either the left or right hand. Adopting a combination of univariate and multivariate analyses, we revealed specific patterns of beta power modulations underlying varying levels of neural representations during movement planning. (1) Effector-specific modulations were evident as a decrease in power in the beta band. Within both hemispheres, this decrease was stronger while planning a movement with the contralateral hand. (2) The comparison of planned grasping and reaching led to a relative increase in power in the beta band. These power changes were localized within temporal, premotor and posterior parietal cortices. Action-related modulations overlapped with effector-related beta power changes within widespread frontal and parietal regions, suggesting the possible integration of these two types of neural representations. (3) Multivariate analyses of action-specific power changes revealed that part of this broadband beta modulation also contributed to the encoding of an effector-independent neural representation of a planned action within fronto-parietal and temporal regions. Our results suggest that beta band power modulations play a central role in movement planning, within both the dorsal and ventral stream, by coding and integrating different levels of neural representations, ranging from the simple representation of the to-be-moved effector up to an abstract, effector-independent representation of the upcoming action.

The origin of word-related motor activity.

Conceptual processing of verbs consistently recruits the left posterior middle temporal gyrus (lpMTG). The left precentral motor cortex also responds to verbs, with higher activity for action than nonaction verbs. The early timing of this effect has suggested that motor features of words' meaning are accessed directly, bypassing access to conceptual representations in lpMTG. An alternative hypothesis is that the retrieval of conceptual representations in lpMTG is necessary to drive more specific, motor-related representations in the precentral gyrus. To test these hypotheses, we first showed that repetitive transcranial magnetic stimulation (rTMS) applied to the verb-preferring lpMTG site selectively impoverished the semantic processing of verbs. In a second experiment, rTMS perturbation of lpMTG, relative to no stimulation (no-rTMS), eliminated the action-nonaction verb distinction in motor activity, as indexed by motor-evoked potentials induced in peripheral muscles with single-pulse TMS over the left primary motor cortex. rTMS pertubation of an occipital control site, relative to no-rTMS, did not affect the action-nonaction verb distinction in motor activity, but the verb contrast did not differ reliably from the lpMTG effect. The results show that lpMTG carries core semantic information necessary to drive the activation of specific (motor) features in the precentral gyrus.

The effect of goals and vision on movements: a case study of optic ataxia and limb apraxia.

Normally we can perform a variety of goal-directed movements effortlessly. However, damage to the parietal cortex may dramatically reduce this ability, giving rise to optic ataxia and limb apraxia. Patients with optic ataxia show clear misreaches towards targets when presented in the peripheral visual field, whereas limb apraxia refers to the inability to use common tools or to imitate simple gestures. In the present paper we describe the case of a left-brain damaged patient, who presented both symptoms. We systematically investigated both spatial and temporal parameters of his movements, when asked to reach and grasp common objects to move (Experiment 1) or to use them (Experiment 2), presented either in the central or peripheral visual field. Different movement parameters changed in relation to the goal of the task (grasp to move vs. grasp to use), reflecting a normal modulation of the movement to accomplish tasks with different goals. On the other hand, grip aperture appeared to be more affected from both task goal and viewing condition, with a specific decrement observed when CF was asked to use objects presented peripherally. On the contrary, a neat effect of the viewing condition was observed in the spatial distribution of the end-points of the movements, and of the horizontal end point in particular, which were shifted towards the fixation point when reaching towards peripheral targets. We hypothesized that optic ataxia and limb apraxia have a differential effect on the patient's performance. The specific presence of optic ataxia would have an effect on the movement trajectory, but both symptoms might interact and influence the grasping component of the movement. As a 'cognitive side of motor control impairment', the presence of limb apraxia may have increased the task demands in grasping to use the objects thus exacerbating optic ataxia.