Search

Previous functional imaging studies demonstrated body-selective patches in the primate visual temporal cortex, comparing activations to static bodies and static images of other categories. However, the use of static instead of dynamic displays of moving bodies may have underestimated the extent of the body patch network. Indeed, body dynamics provide information about action and emotion and may be processed in patches not activated by static images. Thus, to map with fMRI the full extent of the macaque body patch system in the visual temporal cortex, we employed dynamic displays of natural-acting monkey bodies, dynamic monkey faces, objects, and scrambled versions of these videos, all presented during fixation. We found nine body patches in the visual temporal cortex, starting posteriorly in the superior temporal sulcus (STS) and ending anteriorly in the temporal pole. Unlike for static images, body patches were present consistently in both the lower and upper banks of the STS. Overall, body patches showed a higher activation by dynamic displays than by matched static images, which, for identical stimulus displays, was less the case for the neighboring face patches. These data provide the groundwork for future single-unit recording studies to reveal the spatiotemporal features the neurons of these body patches encode. These fMRI findings suggest that dynamics have a stronger contribution to population responses in body than face patches.

The dorsal premotor cortex (PMd) plays an essential role in visually guided goal-directed motor behavior. Although there are several planning processes for achieving goal-directed behavior, the separate neural processes are largely unknown. Here, we created a new visuo-goal task to investigate the step-by-step planning processes for visuomotor and visuo-goal behavior in humans. Using functional magnetic resonance imaging, we found activation in different portions of the bilateral PMd during each processing step. In particular, the activated area for rule-based visuomotor and visuo-goal mapping was located at the ventrorostral portion of the bilateral PMd, that for action plan specification was at the dorsocaudal portion of the left PMd, that for transformation was at the rostral portion of the left PMd, and that for action preparation was at the caudal portion of the bilateral PMd. Thus, the left PMd was involved throughout all of the processes, but the right PMd was involved only in rule-based visuomotor and visuo-goal mapping and action preparation. The locations related to each process were generally spatially separated from each other, but they overlapped partially. These findings revealed that there are functional subregions in the bilateral PMd in humans and these subregions form a functional gradient to achieve goal-directed behavior.

While brain research over the past decades has shed light on the neural correlates of social cognition and behavior in human and non-human primates, most of this research has been performed in virtual settings requiring subjects to observe pictures or recorded videos instead of observing or interacting with another real-live individual. Here we present a two-monkey fMRI setup, allowing examining whole brain responses in macaque monkeys while they observe or interact face-to-face with another real-live conspecific. We tested this setup by comparing overall brain responses during observation of conspecific hand actions in a virtual (observation of recorded videos of actions) or live context (observation of a real-live conspecific performing actions). This dyadic monkey fMRI setup allows examining brain-wide responses in macaque monkeys during different aspects of social behavior, including observation of real-live actions and sensations, social facilitation, joint-attention and social interactions.

While mirror neurons have been found in several monkey brain regions, their existence in the human brain is still largely inferred from indirect non-invasive measurements like functional MRI. It has been proposed that, beyond showing overlapping brain responses during action observation and execution tasks, candidate mirror neuron regions should demonstrate cross-modal action specificity, in line with a defining physiological characteristic of these neurons in the monkey brain. Although cross-modal fMRI adaptation has been put forward as a suited method to test this key feature of cross-modal action specificity in the human brain, so far, the overall usefulness of this technique to demonstrate mirror neuron activity remains unclear. To date, it has never been tested to what extent monkey brain regions known to house mirror neurons, would yield uni- and/or cross-modal fMRI adaptation effects. We therefore performed an fMRI adaptation experiment while male rhesus macaques either performed or observed two different goal-directed hand actions. Executing grasp/lift or touch/press actions in the dark, as well as observing videos of these monkey hand actions, yielded robust responses throughout the brain, including overlapping fMRI responses in parietal and premotor mirror neuron regions. Uni-modal adaptation effects were mostly restricted to the visual modality and the early visual cortices. Both frequentist and Bayesian statistical analyses however suggested no evidence for cross-modal fMRI adaptation effects in monkey parietal and premotor mirror neuron regions. Overall, these findings suggest monkey mirror neuron activity does not readily translate into cross-modal repetition suppression effects that can be detected by fMRI.

To interact meaningfully with its environment, an agent must integrate external information with its own internal states. However, information about the environment is often noisy. In this study, we identify a neural correlate that tracks how asymmetries between competing alternatives evolve over the course of a decision. In our task participants had to monitor a stream of discrete visual stimuli over time and decide whether or not to act, on the basis of either strong or ambiguous evidence. We found that the classic P3 event-related potential evoked by sequential evidence items tracked decision-making processes and predicted participants’ categorical choices on a single trial level, both when evidence was strong and when it was ambiguous. The P3 amplitudes in response to evidence supporting the eventually selected option increased over trial time as decisions evolved, being maximally different from the P3 amplitudes evoked by competing evidence at the time of decision. Computational modelling showed that both the neural dynamics and behavioural primacy and recency effects can be explained by a combination of (a) competition between mutually inhibiting accumulators for the two categorical choice outcomes, and (b) a context-dependant urgency signal. In conditions where evidence was presented at a low rate, urgency increased faster than in conditions when evidence was very frequent. We also found that the readiness potential, a classic marker of endogenously initiated actions, was observed preceding movements in all conditions - even when those were strongly driven by external evidence.

People are capable of rapid improvements in performance when they are offered a reward. The neural mechanism by which this performance enhancement occurs remains unclear. We investigated this phenomenon by offering people monetary reward for successful performance in a sequence production task. We found that people performed actions more quickly and accurately when they were offered large reward. Increasing reward magnitude was associated with elevated activity throughout the brain prior to movement. Multivariate patterns of activity in these reward-responsive regions encoded information about the upcoming action. Follow-up analyses provided evidence that action decoding in pre-SMA and other motor planning areas was improved for large reward trials and successful action decoding in SMA was associated with improved performance. These results suggest that reward may enhance performance by enhancing neural representations of action used in motor planning.

In sensorimotor integration, the brain needs to decide how its predictions should accommodate novel evidence by ‘gating’ sensory data depending on the current context. Here, we examined the oscillatory correlates of this process by recording magnetoencephalography (MEG) data during a new task requiring action under intersensory conflict. We used virtual reality to decouple visual (virtual) and proprioceptive (real) hand postures during a task in which the phase of grasping movements tracked a target (in either modality). Thus, we rendered visual information either task-relevant or a (to-be-ignored) distractor. Under visuo-proprioceptive incongruence, occipital beta power decreased (relative to congruence) when vision was task-relevant but increased when it had to be ignored. Dynamic causal modeling (DCM) revealed that this interaction was best explained by diametrical, task-dependent changes in visual gain. These results suggest a crucial role for beta oscillations in the contextual gating (i.e., gain or precision control) of visual vs proprioceptive action feedback, depending on current behavioral demands.

Recent evidence points to a role of the primary visual cortex that goes beyond visual processing into high-level cognitive and motor-related functions, including action planning, even in absence of feedforward visual information. It has been proposed that, at the neural level, motor imagery is a simulation based on motor representations, and neuroimaging studies have shown overlapping and shared activity patterns for motor imagery and action execution in frontal and parietal cortices. Yet, the role of the early visual cortex in motor imagery remains unclear. Here we used multivoxel pattern analyses on functional magnetic resonance imaging (fMRI) data to examine whether the content of motor imagery and action intention can be reliably decoded from the activity patterns in the retinotopic location of the target object in the early visual cortex. Further, we investigated whether the discrimination between specific actions generalizes across imagined and intended movements. Eighteen right-handed human participants (11 females) imagined or performed delayed hand actions towards a centrally located object composed of a small shape attached on a large shape. Actions consisted of grasping the large or small shape, and reaching to the center of the object. We found that despite comparable fMRI signal amplitude for different planned and imagined movements, activity patterns in the early visual cortex, as well as dorsal premotor and anterior intraparietal cortex, accurately represented action plans and action imagery. However, movement content is similar irrespective of whether actions are actively planned or covertly imagined in parietal but not early visual or premotor cortex, suggesting a generalized motor representation only in regions that are highly specialized in object directed grasping actions and movement goals. In sum, action planning and imagery have overlapping but non identical neural mechanisms in the cortical action network.

A growing literature supports the existence of interactions between emotion and action in the brain, and the central participation of the anterior midcingulate cortex (aMCC) in this regard. In the present functional magnetic resonance imaging study, we sought to investigate the role of self-relevance during such interactions by varying the context in which threating pictures were presented (with guns pointed towards or away from the observer). Participants performed a simple visual detection task following exposure to such stimuli. Except for voxelwise tests, we adopted a Bayesian analysis framework which evaluated evidence for the hypotheses of interest, given the data, in a continuous fashion. Behaviorally, our results demonstrated a valence by context interaction such that there was a tendency of speeding up responses to targets after viewing threat pictures directed towards the participant. In the brain, interaction patterns that paralleled those observed behaviorally were observed most notably in the middle temporal gyrus, supplementary motor area, precentral gyrus, and anterior insula. In these regions, activity was overall greater during threat conditions relative to neutral ones, and this effect was enhanced in the directed towards context. A valence by context interaction was observed in the aMCC too, where we also observed a correlation (across participants) of evoked responses and reaction time data. Taken together, our study revealed the context-sensitive engagement of motor-related areas during emotional perception, thus supporting the idea that emotion and action interact in important ways in the brain.

“Two route” theories of object-related action processing posit different temporal activation profiles of grasp-to-move actions (rapidly evoked based on object structure) versus skilled use actions (more slowly activated based on semantic knowledge). We capitalized on the exquisite temporal resolution and multidimensionality of Event-Related Potentials (ERPs) to directly test this hypothesis. Participants viewed manipulable objects (e.g., calculator) preceded by objects sharing either “grasp”, “use”, or no action attributes (e.g., bar of soap, keyboard, earring, respectively), as well as by action-unrelated but taxonomically-related objects (e.g., abacus); participants judged whether the two objects were related. The results showed more positive responses to “grasp-to-move” primed objects than “skilled use” primed objects or unprimed objects starting in the P1 (0–150 ms) time window and continuing onto the subsequent N1 and P2 components (150–300 ms), suggesting that only “grasp-to-move”, but not “skilled use”, actions may facilitate visual attention to object attributes. Furthermore, reliably reduced N400s (300–500 ms), an index of semantic processing, were observed to taxonomically primed and “skilled use” primed objects relative to unprimed objects, suggesting that “skilled use” action attributes are a component of distributed, multimodal semantic representations of objects. Together, our findings provide evidence supporting two-route theories by demonstrating that “grasp-to-move” and “skilled use” actions impact different aspects of object processing and highlight the relationship of “skilled use” information to other aspects of semantic memory. [ABSTRACT FROM AUTHOR]

• We mapped patches in visual temporal cortex that are activated by dynamic bodies. • The body patch network is more extensive for dynamic than static bodies. • Response to dynamic bodies in upper and lower bank of the superior temporal sulcus. • Stronger effect of dynamics in body patches than in neighboring face patches. Previous functional imaging studies demonstrated body-selective patches in the primate visual temporal cortex, comparing activations to static bodies and static images of other categories. However, the use of static instead of dynamic displays of moving bodies may have underestimated the extent of the body patch network. Indeed, body dynamics provide information about action and emotion and may be processed in patches not activated by static images. Thus, to map with fMRI the full extent of the macaque body patch system in the visual temporal cortex, we employed dynamic displays of natural-acting monkey bodies, dynamic monkey faces, objects, and scrambled versions of these videos, all presented during fixation. We found nine body patches in the visual temporal cortex, starting posteriorly in the superior temporal sulcus (STS) and ending anteriorly in the temporal pole. Unlike for static images, body patches were present consistently in both the lower and upper banks of the STS. Overall, body patches showed a higher activation by dynamic displays than by matched static images, which, for identical stimulus displays, was less the case for the neighboring face patches. These data provide the groundwork for future single-unit recording studies to reveal the spatiotemporal features the neurons of these body patches encode. These fMRI findings suggest that dynamics have a stronger contribution to population responses in body than face patches. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The dorsal premotor cortex (PMd) plays an essential role in visually guided goal-directed motor behavior. Although there are several planning processes for achieving goal-directed behavior, the separate neural processes are largely unknown. Here, we created a new visuo-goal task to investigate the step-by-step planning processes for visuomotor and visuo-goal behavior in humans. Using functional magnetic resonance imaging, we found activation in different portions of the bilateral PMd during each processing step. In particular, the activated area for rule-based visuomotor and visuo-goal mapping was located at the ventrorostral portion of the bilateral PMd, that for action plan specification was at the dorsocaudal portion of the left PMd, that for transformation was at the rostral portion of the left PMd, and that for action preparation was at the caudal portion of the bilateral PMd. Thus, the left PMd was involved throughout all of the processes, but the right PMd was involved only in rule-based visuomotor and visuo-goal mapping and action preparation. The locations related to each process were generally spatially separated from each other, but they overlapped partially. These findings revealed that there are functional subregions in the bilateral PMd in humans and these subregions form a functional gradient to achieve goal-directed behavior. [ABSTRACT FROM AUTHOR]

Previous theoretical and experimental works has shown that preparing to act causes enhanced perceptual processing at movement-relevant locations. Up until now, this has focused almost exclusively on the goal of an action, neglecting the role of the effector. We addressed this by measuring changes in visual processing across time during motor preparation at both goal and effector locations. We compared event related potentials (ERPs) elicited by task-irrelevant visual probe stimuli at both goal and effector locations during motor preparation. Participants were instructed to place their hands on two starting positions (effector locations) and an auditory tone instructed them to immediately move to one of two target buttons (goal locations). Probe stimuli were presented in the interval between the offset of the cue and the execution of the movement at either a goal or an effector location. Probes were presented randomly at either 100 ms, 200 ms or 300 ms after the auditory cue. Analysis of the visual N1 ERP showed enhanced visual processing at moving vs. not-moving goal locations across all three SOAs. At effector locations, enhanced processing for the moving vs. not-moving effector was only observed during the middle (200 ms) SOA. These results demonstrate, for the first time, simultaneous perceptual enhancement of goal and effector locations during motor preparation. We interpret these results as reflecting a temporally and spatially specific dynamic attentional map of the environment that adapts to maximise efficiency of movement by selectively weighting processing of multiple functional components of action in parallel. [ABSTRACT FROM AUTHOR]

While brain research over the past decades has shed light on the neural correlates of social cognition and behavior in human and non-human primates, most of this research has been performed in virtual settings requiring subjects to observe pictures or recorded videos instead of observing or interacting with another real-live individual. Here we present a two-monkey fMRI setup, allowing examining whole brain responses in macaque monkeys while they observe or interact face-to-face with another real-live conspecific. We tested this setup by comparing overall brain responses during observation of conspecific hand actions in a virtual (observation of recorded videos of actions) or live context (observation of a real-live conspecific performing actions). This dyadic monkey fMRI setup allows examining brain-wide responses in macaque monkeys during different aspects of social behavior, including observation of real-live actions and sensations, social facilitation, joint-attention and social interactions. [ABSTRACT FROM AUTHOR]

Abstract: Naturalistic observation and experimental studies in humans and other primates show that observing an individual in need automatically triggers helping behavior. The aim of the present study is to clarify the neurofunctional basis of social influences on individual helping behavior. We investigate whether when participants witness an emergency, while performing an unrelated color-naming task in an fMRI scanner, the number of bystanders present at the emergency influences neural activity in regions related to action preparation. The results show a decrease in activity with the increase in group size in the left pre- and postcentral gyri and left medial frontal gyrus. In contrast, regions related to visual perception and attention show an increase in activity. These results demonstrate the neural mechanisms of social influence on automatic action preparation that is at the core of helping behavior when witnessing an emergency. [Copyright &y& Elsevier]

Abstract: The idea that the conceptual system draws on sensory and motor systems has received considerable experimental support in recent years. Whether the tight coupling between sensory-motor and conceptual systems is modulated by factors such as context or task demands is a matter of controversy. Here, we tested the context sensitivity of this coupling by using action verbs in three different types of sentences in an fMRI study: literal action, apt but non-idiomatic action metaphors, and action idioms. Abstract sentences served as a baseline. The result showed involvement of sensory-motor areas for literal and metaphoric action sentences, but not for idiomatic ones. A trend of increasing sensory-motor activation from abstract to idiomatic to metaphoric to literal sentences was seen. These results support a gradual abstraction process whereby the reliance on sensory-motor systems is reduced as the abstractness of meaning as well as conventionalization is increased, highlighting the context sensitive nature of semantic processing. [Copyright &y& Elsevier]

Abstract: Decision-making invokes two fundamental axes of control: affect or valence, spanning reward and punishment, and effect or action, spanning invigoration and inhibition. We studied the acquisition of instrumental responding in healthy human volunteers in a task in which we orthogonalized action requirements and outcome valence. Subjects were much more successful in learning active choices in rewarded conditions, and passive choices in punished conditions. Using computational reinforcement-learning models, we teased apart contributions from putatively instrumental and Pavlovian components in the generation of the observed asymmetry during learning. Moreover, using model-based fMRI, we showed that BOLD signals in striatum and substantia nigra/ventral tegmental area (SN/VTA) correlated with instrumentally learnt action values, but with opposite signs for go and no-go choices. Finally, we showed that successful instrumental learning depends on engagement of bilateral inferior frontal gyrus. Our behavioral and computational data showed that instrumental learning is contingent on overcoming inherent and plastic Pavlovian biases, while our neuronal data showed this learning is linked to unique patterns of brain activity in regions implicated in action and inhibition respectively. [Copyright &y& Elsevier]

Abstract: When we observe an action, we recognize meaningful action steps that help us to predict probable upcoming action steps. This segmentation of observed actions, or more generally events, has been proposed to rely in part on changes in motion features. However, segmentation of actions, in contrast to meaningless movements, may exploit additional information such as action knowledge. The present fMRI study sought to tear apart the neural signatures of processing two sources of information that observers may exploit at action boundaries: change in motion dynamics and action knowledge. To this end, subjects performed a segmentation task on both actions (that can be segmented based on motion and action knowledge) as well as tai chi movements (that can be segmented only based on motion) and two further control conditions that implemented point-light walker like displays of the same videos. Behavioral tests showed that motion features played a critical role in boundary detection in all conditions. Consistent with this finding, activity in area MT was enhanced during boundary detection in all conditions, but importantly, this effect was not stronger for actions. In contrast, only action boundary detection was reflected by specific activation in the superior frontal sulcus, parietal angular gyrus and the parahippocampal cortex. Based on these findings, we propose that during action observation, motion features trigger a top–down modulation of the attentional focus and the incitement of retrieving long-term memory place-action associations. While action perception entails activity common to processing of all motion stimuli, it is at the same time unique as it allows long-term memory based predictions of succeeding steps. [Copyright &y& Elsevier]

Abstract: The organization of the cerebral cortex into distinct modules may be described along several dimensions, most importantly, structure, connectivity and function. Identification of cortical modules by differences in whole-brain connectivity profiles derived from diffusion tensor imaging or resting state correlations has already been shown. These approaches, however, carry no task-related information. Hence, inference on the functional relevance of the ensuing parcellation remains tentative. Here, we demonstrate, that Meta-Analytic Connectivity Modeling (MACM) allows the delineation of cortical modules based on their whole-brain co-activation pattern across databased neuroimaging results. Using a model free approach, two regions of the medial pre-motor cortex, SMA and pre-SMA were differentiated solely based on their functional connectivity. Assessing the behavioral domain and paradigm class meta-data of the experiments associated with the clusters derived from the co-activation based parcellation moreover allows the identification of their functional characteristics. The ensuing hypotheses about functional differentiation and distinct functional connectivity between pre-SMA and SMA were then explicitly tested and confirmed in independent datasets using functional and resting state fMRI. Co-activation based parcellation thus provides a new perspective for identifying modules of functional connectivity and linking them to functional properties, hereby generating new and subsequently testable hypotheses about the organization of cortical modules. [Copyright &y& Elsevier]

Abstract: Embodied accounts of language processing suggest that sensorimotor areas, generally dedicated to perception and action, are also involved in the processing and representation of word meaning. Support for such accounts comes from studies showing that language about actions selectively modulates the execution of congruent and incongruent motor responses (e.g., Glenberg & Kaschak, 2002), and from functional neuroimaging studies showing that understanding action-related language recruits sensorimotor brain areas (e.g. Hauk, Johnsrude, & Pulvermueller, 2004). In the current experiment we explored the basis of the neural motor system''s involvement in representing words denoting actions. Specifically, we investigated whether the motor system''s involvement is modulated by the specificity of the kinematics associated with a word. Previous research in the visual domain indicates that words denoting basic level category members lacking a specific form (e.g., bird) are less richly encoded within visual areas than words denoting subordinate level members (e.g., pelican), for which the visual form is better specified (Gauthier, Anderson, Tarr, Skudlarski, & Gore, 1997). In the present study we extend these findings to the motor domain. Modulation of the BOLD response elicited by verbs denoting a general motor program (e.g., to clean) was compared to modulation elicited by verbs denoting a more specific motor program (e.g., to wipe). Conform with our hypothesis, a region within the bilateral inferior parietal lobule, typically serving the representation of action plans and goals, was sensitive to the specificity of motor programs associated with the action verbs. These findings contribute to the growing body of research on embodied language representations by showing that the concreteness of an action-semantic feature is reflected in the neural response to action verbs. [Copyright &y& Elsevier]

Abstract: Sensorimotor-based theories of semantic memory contend that semantic information about an object is represented in the neural substrate invoked when we perceive or interact with it. We used fMRI adaptation to test this prediction, measuring brain activation as participants read pairs of words. Pairs shared function (flashlight–lantern), shape (marble–grape), both (pencil–pen), were unrelated (saucer–needle), or were identical (drill–drill). We observed adaptation for pairs with both function and shape similarity in left premotor cortex. Further, degree of function similarity was correlated with adaptation in three regions: two in the left temporal lobe (left medial temporal lobe, left middle temporal gyrus), which has been hypothesized to play a role in mutimodal integration, and one in left superior frontal gyrus. We also found that degree of manipulation (i.e., action) and function similarity were both correlated with adaptation in two regions: left premotor cortex and left intraparietal sulcus (involved in guiding actions). Additional considerations suggest that the adaptation in these two regions was driven by manipulation similarity alone; thus, these results imply that manipulation information about objects is encoded in brain regions involved in performing or guiding actions. Unexpectedly, these same two regions showed increased activation (rather than adaptation) for objects similar in shape. Overall, we found evidence (in the form of adaptation) that objects that share semantic features have overlapping representations. Further, the particular regions of overlap provide support for the existence of both sensorimotor and amodal/multimodal representations. [Copyright &y& Elsevier]

Abstract: Action representations can interact with object recognition processes. For example, so-called mirror neurons respond both when performing an action and when seeing or hearing such actions. Investigations of auditory object processing have largely focused on categorical discrimination, which begins within the initial 100 ms post-stimulus onset and subsequently engages distinct cortical networks. Whether action representations themselves contribute to auditory object recognition and the precise kinds of actions recruiting the auditory–visual mirror neuron system remain poorly understood. We applied electrical neuroimaging analyses to auditory evoked potentials (AEPs) in response to sounds of man-made objects that were further subdivided between sounds conveying a socio-functional context and typically cuing a responsive action by the listener (e.g. a ringing telephone) and those that are not linked to such a context and do not typically elicit responsive actions (e.g. notes on a piano). This distinction was validated psychophysically by a separate cohort of listeners. Beginning ~300 ms, responses to such context-related sounds significantly differed from context-free sounds both in the strength and topography of the electric field. This latency is >200 ms subsequent to general categorical discrimination. Additionally, such topographic differences indicate that sounds of different action sub-types engage distinct configurations of intracranial generators. Statistical analysis of source estimations identified differential activity within premotor and inferior (pre)frontal regions (Brodmann''s areas (BA) 6, BA8, and BA45/46/47) in response to sounds of actions typically cuing a responsive action. We discuss our results in terms of a spatio-temporal model of auditory object processing and the interplay between semantic and action representations. [Copyright &y& Elsevier]

Abstract: Several studies indicate that both posterior superior temporal sulcus/middle temporal gyrus (pSTS/MTG) and left inferior frontal gyrus (LIFG) are involved in integrating information from different modalities. Here we investigated the respective roles of these two areas in integration of action and language information. We exploited the fact that the semantic relationship between language and different forms of action (i.e. co-speech gestures and pantomimes) is radically different. Speech and co-speech gestures are always produced together, and gestures are not unambiguously understood without speech. On the contrary, pantomimes are not necessarily produced together with speech and can be easily understood without speech. We presented speech together with these two types of communicative hand actions in matching or mismatching combinations to manipulate semantic integration load. Left and right pSTS/MTG were only involved in semantic integration of speech and pantomimes. Left IFG on the other hand was involved in integration of speech and co-speech gestures as well as of speech and pantomimes. Effective connectivity analyses showed that depending upon the semantic relationship between language and action, LIFG modulates activation levels in left pSTS. This suggests that integration in pSTS/MTG involves the matching of two input streams for which there is a relatively stable common object representation, whereas integration in LIFG is better characterized as the on-line construction of a new and unified representation of the input streams. In conclusion, pSTS/MTG and LIFG are differentially involved in multimodal integration, crucially depending upon the semantic relationship between the input streams. [Copyright &y& Elsevier]

Abstract: Being exposed to fear or anger signals makes us feel threatened and prompts us to prepare an adaptive response. Yet, while fear and anger behaviors are both threat signals, what counts as an adaptive response is often quite different. In contrast with fear, anger is often displayed with the aim of altering the behavior of the agent to which it is addressed. To identify brain responses that are common or specific to the perception of these two types of threat signals, we used functional magnetic resonance imaging and asked subjects to recognize dynamic actions expressing fear, anger and neutral behaviors. As compared with neutral actions, the perception of fear and anger behaviors elicited comparable activity increases in the left amygdala and temporal cortices as well as in the ventrolateral and the dorsomedial prefrontal cortex. Whereas the perception of fear elicited specific activity in the right temporoparietal junction, the perception of anger triggered condition-specific activity in a wider set of regions comprising the anterior temporal lobe, the premotor cortex and the ventromedial prefrontal cortex, consistent with the hypothesis that coping with threat from exposure to anger requires additional contextual information and behavioral adjustments. [Copyright &y& Elsevier]

Abstract: This note presents a simple Bayesian filtering scheme, using variational calculus, for inference on the hidden states of dynamic systems. Variational filtering is a stochastic scheme that propagates particles over a changing variational energy landscape, such that their sample density approximates the conditional density of hidden and states and inputs. The key innovation, on which variational filtering rests, is a formulation in generalised coordinates of motion. This renders the scheme much simpler and more versatile than existing approaches, such as those based on particle filtering. We demonstrate variational filtering using simulated and real data from hemodynamic systems studied in neuroimaging and provide comparative evaluations using particle filtering and the fixed-form homologue of variational filtering, namely dynamic expectation maximisation. [Copyright &y& Elsevier]

Abstract: This paper presents a variational treatment of dynamic models that furnishes time-dependent conditional densities on the path or trajectory of a system''s states and the time-independent densities of its parameters. These are obtained by maximising a variational action with respect to conditional densities, under a fixed-form assumption about their form. The action or path-integral of free-energy represents a lower bound on the model''s log-evidence or marginal likelihood required for model selection and averaging. This approach rests on formulating the optimisation dynamically, in generalised coordinates of motion. The resulting scheme can be used for online Bayesian inversion of nonlinear dynamic causal models and is shown to outperform existing approaches, such as Kalman and particle filtering. Furthermore, it provides for dual and triple inferences on a system''s states, parameters and hyperparameters using exactly the same principles. We refer to this approach as dynamic expectation maximisation (DEM). [Copyright &y& Elsevier]

Abstract: Both animal and human studies suggest that the efficiency with which we are able to grasp objects is attributable to a repertoire of motor signals derived directly from vision. This is in general agreement with the long-held belief that the automatic generation of motor signals by the perception of objects is based on the actions they afford. In this study, we used magnetoencephalography (MEG) to determine the spatial distribution and temporal dynamics of brain regions activated during passive viewing of object and non-object targets that varied in the extent to which they afforded a grasping action. Synthetic Aperture Magnetometry (SAM) was used to localize task-related oscillatory power changes within specific frequency bands, and the time course of activity within given regions-of-interest was determined by calculating time–frequency plots using a Morlet wavelet transform. Both single subject and group-averaged data on the spatial distribution of brain activity are presented. We show that: (i) significant reductions in 10–25 Hz activity within extrastriate cortex, occipito-temporal cortex, sensori-motor cortex and cerebellum were evident with passive viewing of both objects and non-objects; and (ii) reductions in oscillatory activity within the posterior part of the superior parietal cortex (area Ba7) were only evident with the perception of objects. Assuming that focal reductions in low-frequency oscillations (<30 Hz) reflect areas of heightened neural activity, we conclude that: (i) activity within a network of brain areas, including the sensori-motor cortex, is not critically dependent on stimulus type and may reflect general changes in visual attention; and (ii) the posterior part of the superior parietal cortex, area Ba7, is activated preferentially by objects and may play a role in computations related to grasping. [Copyright &y& Elsevier]

Abstract: Many studies have reported a movement-related modulation of response in the primary and secondary somatosensory cortices (SI and SII) to a task-irrelevant stimulation in primates. In the present study, magnetoencephalography (MEG) was used to examine the top–down centrifugal regulation of neural responses in the human SI and SII to a task-relevant somatosensory signal triggering a voluntary movement. Nine healthy adults participated in the study. A visual warning signal was followed 2 s later by a somatosensory imperative signal delivered to the right median nerve at the wrist. Three kinds of warning signal informed the participants of the reaction which should be executed on presentation of the imperative signal (rest or extension of the right index finger, extension of the left index finger). The somatosensory stimulation was used to both generate neural responses and trigger voluntary movement and therefore was regarded as a task-relevant signal. The responses were recorded using a whole-head MEG system. The P35m response around the SI was reduced in magnitude without alteration of the primary SI response, N20m, when the signal triggered a voluntary movement compared to the control condition, whereas bilateral SII responses peaking at 70–100 ms were enhanced and the peak latency was shortened. The peak latency of the responses in the SI and SII preceded the onset of the earliest voluntary muscle activation in each subject. Later bilateral perisylvian responses were also enhanced with movement. In conclusion, neural activities in the SI and SII evoked by task-relevant somatosensory signals are regulated differently by motor-related neural activities before the afferent inputs. The present findings indicate a difference in function between the SI and SII in somatosensory–motor regulation. [Copyright &y& Elsevier]

Abstract: Human behavior is often dictated by rules or prescribed guides for action. Little is currently known regarding how these rules are stored in long-term memory or retrieved and implemented. Here, we examined the roles of ventrolateral prefrontal cortex (VLPFC) and posterior middle temporal gyrus (postMTG) in rule use. We tested two hypotheses: first, that knowledge about actions associated with abstract visual symbols is stored in postMTG, and second, that VLPFC is involved in the controlled retrieval of rule meanings. Subjects viewed a series of road signs during event-related fMRI data collection. Three types of signs were intermixed: highly familiar signs, novel signs whose meaning was explained to subjects prior to scanning, and novel signs whose meaning was not explained. Subjects were asked to think about the meaning of each sign as it was presented during scanning and then to give its meaning in a post-scan test. Left postMTG was more active when subjects viewed signs whose meaning they knew than signs whose meaning they did not know, consistent with a role in storing rule meanings. This region was not modulated by experience, in that it was equally engaged by newly trained and well-learned signs. In contrast, right VLPFC was more active for newly trained signs than for either well-learned or incorrect ones, consistent with a role in controlled retrieval. Left VLPFC was reliably engaged while subjects attempted to interpret the signs but did not differ according to knowledge or experience. These data implicate postMTG in rule storage and VLPFC in rule retrieval. [Copyright &y& Elsevier]

Engagement of the primary motor cortex (MI) during the observation of actions has been debated for a long time. In the present study, we used the quantitative 14C-deoxyglucose method in monkeys that either grasped 3-D objects or observed the same movements executed by humans. We found that the forelimb regions of the MI and the primary somatosensory (SI) cortex were significantly activated in both cases. Our study resolves a debate in the literature, providing strong evidence for use of MI representations during the observation of actions. It demonstrates that the observation of an action is represented in the primary motor and somatosensory cortices as is its execution. It indicates that in terms of neural correlates, recognizing a motor behavior is like executing the same behavior, requiring the involvement of a distributed system encompassing not only the premotor but also the primary motor cortex. We suggest that movements and their proprioceptive components are stored as motor and somatosensory representations in motor and somatosensory cortices, respectively, and that these representations are recalled during observation of an action. [Copyright &y& Elsevier]

Actions are said to be ‘willed’ if we consciously pay attention to their selection. It has been suggested that they are associated with activations in the dorsal prefrontal cortex (area 46). However, because previous experiments typically used a ‘free selection’ paradigm to examine this hypothesis, it is unclear whether the results reflected the attention to the selection of action or the freedom of choice allowed by the tasks. In this experiment, we minimized the difference of working memory demand across task conditions by using novel stimuli in each trial. We found that activation in the dorsal prefrontal cortex on a free selection task was not significantly different from that induced by another task that required attention to the selection of action, although the responses were externally specified. This suggests that the dorsal prefrontal cortex is in fact associated with attention to the selection of action, but does not play a unique role in the generation of internally initiated actions. However, the presupplementary motor area (pre-SMA) may subserve this function as activity in this region was found to be tightly associated with the free selection of responses. [Copyright &y& Elsevier]

Imitation is a natural mechanism involving perception–action coupling which plays a foundational role in human development, in particular to extract the intention from the surface behavior exhibited by others. The aim of this H152O PET activation experiment was to investigate the neural basis of imitation of object-oriented actions in normal adults. Experimental conditions were derived from a factorial design. The factors were: (a) is the stimulus event shown to subjects during observation of the model and (b) is the response manipulation performed by the subject. Two key components of human action, the goal and the means to achieve it, were systematically investigated. The results revealed partially overlapping clusters of increased regional cerebral blood flow in the right dorsolateral prefrontal area and in cerebellum when subjects imitated either of the two components. Moreover, specific activity was detected in the medial prefrontal cortex during the imitation of the means, whereas imitating the goal was associated with increased activity in the left premotor cortex. Our results suggest that for normally functioning adults, imitating a gesture activates neural processing of the intention (or goal) underlying the observed action. [Copyright &y& Elsevier]

While mirror neurons have been found in several monkey brain regions, their existence in the human brain is still largely inferred from indirect non-invasive measurements like functional MRI. It has been proposed that, beyond showing overlapping brain responses during action observation and execution tasks, candidate mirror neuron regions should demonstrate cross-modal action specificity, in line with a defining physiological characteristic of these neurons in the monkey brain. Although cross-modal fMRI adaptation has been put forward as a suited method to test this key feature of cross-modal action specificity in the human brain, so far, the overall usefulness of this technique to demonstrate mirror neuron activity remains unclear. To date, it has never been tested to what extent monkey brain regions known to house mirror neurons, would yield uni- and/or cross-modal fMRI adaptation effects. We therefore performed an fMRI adaptation experiment while male rhesus macaques either performed or observed two different goal-directed hand actions. Executing grasp/lift or touch/press actions in the dark, as well as observing videos of these monkey hand actions, yielded robust responses throughout the brain, including overlapping fMRI responses in parietal and premotor mirror neuron regions. Uni-modal adaptation effects were mostly restricted to the visual modality and the early visual cortices. Both frequentist and Bayesian statistical analyses however suggested no evidence for cross-modal fMRI adaptation effects in monkey parietal and premotor mirror neuron regions. Overall, these findings suggest monkey mirror neuron activity does not readily translate into cross-modal repetition suppression effects that can be detected by fMRI. [ABSTRACT FROM AUTHOR]

• The classic P3 tracks the evolution of an internal category-specific decision variable in humans. • The amplitude of the P3 evoked by competing evidence items predicts choice on a single-trial level. • The decision variable is modulated by an internal time-dependant urgency signal. • This urgency signal was stronger in conditions with lower rates of evidence. • Primacy and recency effects can be accounted for by context-dependant urgency modulations. To interact meaningfully with its environment, an agent must integrate external information with its own internal states. However, information about the environment is often noisy. In this study, we identify a neural correlate that tracks how asymmetries between competing alternatives evolve over the course of a decision. In our task participants had to monitor a stream of discrete visual stimuli over time and decide whether or not to act, on the basis of either strong or ambiguous evidence. We found that the classic P3 event-related potential evoked by sequential evidence items tracked decision-making processes and predicted participants' categorical choices on a single trial level, both when evidence was strong and when it was ambiguous. The P3 amplitudes in response to evidence supporting the eventually selected option increased over trial time as decisions evolved, being maximally different from the P3 amplitudes evoked by competing evidence at the time of decision. Computational modelling showed that both the neural dynamics and behavioural primacy and recency effects can be explained by a combination of (a) competition between mutually inhibiting accumulators for the two categorical choice outcomes, and (b) a context-dependant urgency signal. In conditions where evidence was presented at a low rate, urgency increased faster than in conditions when evidence was very frequent. We also found that the readiness potential, a classic marker of endogenously initiated actions, was observed preceding movements in all conditions - even when those were strongly driven by external evidence. [ABSTRACT FROM AUTHOR]

• Reward enhanced behavioral performance. • Reward enhanced action decoding in motor planning areas prior to movement. • Action decoding in SMA coincided with improved behavioral performance. • Reward may enhance performance by enhancing action codes used in planning. People are capable of rapid improvements in performance when they are offered a reward. The neural mechanism by which this performance enhancement occurs remains unclear. We investigated this phenomenon by offering people monetary reward for successful performance in a sequence production task. We found that people performed actions more quickly and accurately when they were offered large reward. Increasing reward magnitude was associated with elevated activity throughout the brain prior to movement. Multivariate patterns of activity in these reward-responsive regions encoded information about the upcoming action. Follow-up analyses provided evidence that action decoding in pre-SMA and other motor planning areas was improved for large reward trials and successful action decoding in SMA was associated with improved performance. These results suggest that reward may enhance performance by enhancing neural representations of action used in motor planning. [ABSTRACT FROM AUTHOR]

• We decouple seen and felt hand postures during action via virtual reality. • Vision of the hand is either task-relevant or a distractor. • Task-relevance of vision is reflected by in- or decreases of occipital beta power. • DCM suggests underlying changes in cortical (visual) excitability. • Occipital beta may indicate the contextual gating of visual action feedback. In sensorimotor integration, the brain needs to decide how its predictions should accommodate novel evidence by 'gating' sensory data depending on the current context. Here, we examined the oscillatory correlates of this process by recording magnetoencephalography (MEG) data during a new task requiring action under intersensory conflict. We used virtual reality to decouple visual (virtual) and proprioceptive (real) hand postures during a task in which the phase of grasping movements tracked a target (in either modality). Thus, we rendered visual information either task-relevant or a (to-be-ignored) distractor. Under visuo-proprioceptive incongruence, occipital beta power decreased (relative to congruence) when vision was task-relevant but increased when it had to be ignored. Dynamic causal modeling (DCM) revealed that this interaction was best explained by diametrical, task-dependent changes in visual gain. These results suggest a crucial role for beta oscillations in the contextual gating (i.e., gain or precision control) of visual vs proprioceptive action feedback, depending on current behavioral demands. Image, graphical abstract [ABSTRACT FROM AUTHOR]

Recent evidence points to a role of the primary visual cortex that goes beyond visual processing into high-level cognitive and motor-related functions, including action planning, even in absence of feedforward visual information. It has been proposed that, at the neural level, motor imagery is a simulation based on motor representations, and neuroimaging studies have shown overlapping and shared activity patterns for motor imagery and action execution in frontal and parietal cortices. Yet, the role of the early visual cortex in motor imagery remains unclear. Here we used multivoxel pattern analyses on functional magnetic resonance imaging (fMRI) data to examine whether the content of motor imagery and action intention can be reliably decoded from the activity patterns in the retinotopic location of the target object in the early visual cortex. Further, we investigated whether the discrimination between specific actions generalizes across imagined and intended movements. Eighteen right-handed human participants (11 females) imagined or performed delayed hand actions towards a centrally located object composed of a small shape attached on a large shape. Actions consisted of grasping the large or small shape, and reaching to the center of the object. We found that despite comparable fMRI signal amplitude for different planned and imagined movements, activity patterns in the early visual cortex, as well as dorsal premotor and anterior intraparietal cortex, accurately represented action plans and action imagery. However, movement content is similar irrespective of whether actions are actively planned or covertly imagined in parietal but not early visual or premotor cortex, suggesting a generalized motor representation only in regions that are highly specialized in object directed grasping actions and movement goals. In sum, action planning and imagery have overlapping but non identical neural mechanisms in the cortical action network. • Evidence shows that the early visual cortex (EVC) plays a role in action planning. • Is representation of action intention in EVC related to motor imagery? • Results show that action planning and motor imagery content can be decoded in EVC. • However, action content in EVC is not generalized across planning and imagining tasks. • Parietal cortex shows a generalized representation of action regardless of the task. [ABSTRACT FROM AUTHOR]

A growing literature supports the existence of interactions between emotion and action in the brain, and the central participation of the anterior midcingulate cortex (aMCC) in this regard. In the present functional magnetic resonance imaging study, we sought to investigate the role of self-relevance during such interactions by varying the context in which threating pictures were presented (with guns pointed towards or away from the observer). Participants performed a simple visual detection task following exposure to such stimuli. Except for voxelwise tests, we adopted a Bayesian analysis framework which evaluated evidence for the hypotheses of interest, given the data, in a continuous fashion. Behaviorally, our results demonstrated a valence by context interaction such that there was a tendency of speeding up responses to targets after viewing threat pictures directed towards the participant. In the brain, interaction patterns that paralleled those observed behaviorally were observed most notably in the middle temporal gyrus, supplementary motor area, precentral gyrus, and anterior insula. In these regions, activity was overall greater during threat conditions relative to neutral ones, and this effect was enhanced in the directed towards context. A valence by context interaction was observed in the aMCC too, where we also observed a correlation (across participants) of evoked responses and reaction time data. Taken together, our study revealed the context-sensitive engagement of motor-related areas during emotional perception, thus supporting the idea that emotion and action interact in important ways in the brain. • Increasing threat self-relevance engages motor-related processing areas. • Anterior midcingulate cortex is a node of emotion and motor interaction. • Emotional behavioral modulation is correlated with anterior midcingulate activity. [ABSTRACT FROM AUTHOR]