Nine-month-old infants update their predictive models of a changing environment.

Humans generate internal models of their environment to predict events in the world. As the environments change, our brains adjust to these changes by updating their internal models. Here, we investigated whether and how 9-month-old infants differentially update their models to represent a dynamic environment. Infants observed a predictable sequence of stimuli, which were interrupted by two types of cues. Following the update cue, the pattern was altered, thus, infants were expected to update their predictions for the upcoming stimuli. Because the pattern remained the same after the no-update cue, no subsequent updating was required. Infants showed an amplified negative central (Nc) response when the predictable sequence was interrupted. Late components such as the PSW were also evoked in response to unexpected stimuli; however, we found no evidence for a differential response to the informational value of surprising cues at later stages of processing. Infants rather learned that surprising cues always signal a change in the environment that requires updating. Interestingly, infants responded with an amplified neural response to the absence of an expected change, suggesting a top-down modulation of early sensory processing in infants. Our findings corroborate emerging evidence showing that infants build predictive models early in life.

Infants differentially update their internal models of a dynamic environment.

Unexpected events provide us with opportunities for learning about what to expect from the world around us. Using a saccadic-planning paradigm, we investigated whether and how infants and adults represent the statistics of a changing environment (i.e. build an internal model of the environment). Participants observed differently colored bees that appeared at an unexpected location every few trials. The color cues indicated whether the subsequent bees would appear at this new location (i.e. update trials) or at the same location as previously (i.e. no-update trials). Infants learned the predictive value of the color cues and updated their internal models when necessary. Unlike infants, adults had a tendency to update their models each time they observed a change in the structure. We argue that infants are open to learning from current evidence due to being less influenced by their prior knowledge. This is an advantageous learning strategy to form accurate representations in dynamic environments, which is fundamental for successful adaptation.

How preschoolers and adults represent their joint action partner's behavior.

We investigated the cognitive mechanisms underlying turn-taking joint action in 42-month-old children (Experiment 1) and adults (Experiment 2) using a behavioral task of dressing a virtual bear together. We aimed to investigate how participants represent a partners' behavior, i.e., in terms of specific action kinematics or of action effects. The bear was dressed by pressing a smaller and a bigger button. In the Action-response task, instructions asked participants to respond to the partner by pressing the same or opposite button; in the Action-effect task they had to respond to the partner's action effect by dressing the bear with the lacking part of the clothing, which in some cases implied pressing the same button and in other cases implied pressing the opposite button. In 50% of the trials, the partner's association between each button and the ensuing effect (dressing the bear with t-shirt or pants) was reversed, while it never changed for participants. Both children and adults showed no effect of physical congruency of actions, but showed impaired performance in the Action-effect task if their partner achieved her effect through a different action-effect association than their own. These results suggest that, when encoding their partner's actions, agents are influenced by action-effect associations that they learnt through their own experience. While interference led to overt errors in children, it caused longer reaction times in adults, suggesting that a flexible cognitive control (that is still in development in young children) is required to take on the partner's perspective.

Measuring mimicry: general corticospinal facilitation during observation of naturalistic behaviour.

Mimicry of others' postures and behaviours forms an implicit yet indispensable component of social interactions. However, whereas numerous behavioural studies have investigated the occurrence of mimicry and its social sensitivity, the underlying neurocognitive mechanisms remain elusive. In this study, single-pulse transcranial magnetic stimulation was used to measure corticospinal facilitation during a naturalistic behaviour observation task adapted from the behavioural mimicry literature. Motor evoked potentials (MEPs) in participants' right hands were measured as they observed stimulus videos of a confederate describing photographs. MEPs were recorded while confederates were and were not carrying out hand and leg behaviours that also differed in spatial extent (i.e. large behaviours: face rubbing and leg crossing; small behaviours: finger tapping and foot bouncing). Importantly, the cover task instructions did not refer to the behaviours but instead required participants to focus on the confederates' photograph descriptions in order to later perform a recognition test. A general arousal effect was found, with higher MEPs during stimulus video observation than during a fixation-cross baseline, regardless of whether or not the confederate was carrying out a behaviour at the time of the pulse. When controlling for this general arousal effect, results showed that MEPs during observation of the larger two behaviours were significantly higher than the smaller two behaviours, irrespective of effector. Thus, using a controlled yet naturalistic paradigm, this study suggests that general sensorimotor arousal during social interactions could play a role in implicit behavioural mimicry.

Unravelling the contributions of motor experience and conceptual knowledge in action perception: A training study.

Prior knowledge affects how we perceive the world and the sensorimotor system actively guides our perception. An ongoing dispute regards the extent to which prior motor knowledge versus conceptual knowledge modulates the observation of others' actions. Research indicates that motor experience increases motor activation during action perception. Other research, however, has shown that conceptual familiarity with actions also modulates motor activation, i.e., increased motor activation during observation of unfamiliar, compared to conceptually familiar, actions. To begin to disentangle motor from conceptual contributions to action perception, we uniquely combined motoric and conceptual interventions into one design. We experimentally manipulated participants' experience with both motoric skills and conceptual knowledge, via motor training of kinematically challenging actions and contextual information about the action, respectively, in a week-long training session. Measurements of the effects on motor activity measured via electroencephalography (EEG) during pre- and post-training action observation were compared. We found distinct, non-interacting effects of both manipulations: Motor training increased motor activation, whereas additional conceptual knowledge decreased motor activation. The findings indicate that both factors influence action perception in a distinct and parallel manner. This research speaks to previously irreconcilable findings and provides novel insights about the distinct roles of motor and conceptual contributions to action perception.

Neural mirroring and social interaction: Motor system involvement during action observation relates to early peer cooperation.

Whether we hand over objects to someone, play a team sport, or make music together, social interaction often involves interpersonal action coordination, both during instances of cooperation and entrainment. Neural mirroring is thought to play a crucial role in processing other's actions and is therefore considered important for social interaction. Still, to date, it is unknown whether interindividual differences in neural mirroring play a role in interpersonal coordination during different instances of social interaction. A relation between neural mirroring and interpersonal coordination has particularly relevant implications for early childhood, since successful early interaction with peers is predictive of a more favorable social development. We examined the relation between neural mirroring and children's interpersonal coordination during peer interaction using EEG and longitudinal behavioral data. Results showed that 4-year-old children with higher levels of motor system involvement during action observation (as indicated by lower beta-power) were more successful in early peer cooperation. This is the first evidence for a relation between motor system involvement during action observation and interpersonal coordination during other instances of social interaction. The findings suggest that interindividual differences in neural mirroring are related to interpersonal coordination and thus successful social interaction.

The role of action prediction and inhibitory control for joint action coordination in toddlers.

From early in life, young children eagerly engage in social interactions. Yet, they still have difficulties in performing well-coordinated joint actions with others. Adult literature suggests that two processes are important for smooth joint action coordination: action prediction and inhibitory control. The aim of the current study was to disentangle the potential role of these processes in the early development of joint action coordination. Using a simple turn-taking game, we assessed 2½-year-old toddlers' joint action coordination, focusing on timing variability and turn-taking accuracy. In two additional tasks, we examined their action prediction capabilities with an eye-tracking paradigm and examined their inhibitory control capabilities with a classic executive functioning task (gift delay task). We found that individual differences in action prediction and inhibitory action control were distinctly related to the two aspects of joint action coordination. Toddlers who showed more precision in their action predictions were less variable in their action timing during the joint play. Furthermore, toddlers who showed more inhibitory control in an individual context were more accurate in their turn-taking performance during the joint action. On the other hand, no relation between timing variability and inhibitory control or between turn-taking accuracy and action prediction was found. The current results highlight the distinct role of action prediction and inhibitory action control for the quality of joint action coordination in toddlers. Underlying neurocognitive mechanisms and implications for processes involved in joint action coordination in general are discussed.

Pouring or chilling a bottle of wine: an fMRI study on the prospective planning of object-directed actions.

This fMRI study investigates the neural mechanisms supporting the retrieval of action semantics. A novel motor imagery task was used in which participants were required to imagine planning actions with a familiar object (e.g. a toothbrush) or with an unfamiliar object (e.g. a pair of pliers) based on either goal-related information (i.e. where to move the object) or grip-related information (i.e. how to grasp the object). Planning actions with unfamiliar compared to familiar objects was slower and was associated with increased activation in the bilateral superior parietal lobe, the right inferior parietal lobe and the right insula. The stronger activation in parietal areas for unfamiliar objects fits well with the idea that parietal areas are involved in motor imagery and suggests that this process takes more effort in the case of novel or unfamiliar actions. In contrast, the planning of familiar actions resulted in increased activation in the anterior prefrontal cortex, suggesting that subjects maintained a stronger goal-representation when planning actions with familiar compared to unfamiliar objects. These findings provide further insight into the neural structures that support action semantic knowledge for the functional use of real-world objects and suggest that action semantic knowledge is activated most readily when actions are planned in a goal-directed manner.

Imitation of hand and tool actions is effector-independent.

Following the theoretical notion that tools often extend one's body, in the present study, we investigated whether imitation of hand or tool actions is modulated by effector-specific information. Subjects performed grasping actions toward an object with either a handheld tool or their right hand. Actions were initiated in response to pictures representing a grip at an object that could be congruent or incongruent with the required action (grip-type congruency). Importantly, actions could be cued by means of a tool cue, a hand cue, and a symbolic cue (effector-type congruency). For both hand and tool actions, an action congruency effect was observed, reflected in faster reaction times if the observed grip type was congruent with the required movement. However, neither hand actions nor tool actions were differentially affected by the effector represented in the picture (i.e., when performing a tool action, the action congruency effect was similar for tool cues and hand cues). This finding suggests that imitation of hand and tool actions is effector-independent and thereby supports generalist rather than specialist theories of imitation.

Learning to use novel objects: a training study on the acquisition of novel action representations.

Many studies have suggested that the motor system is organized in a hierarchical fashion, around the prototypical end location associated with using objects. However, most studies supporting the hierarchical view have used well-known actions and objects that are highly over-learned. Accordingly, at present it is unclear if the hierarchical principle applies to learning the use of novel objects as well. In the present study we found that when learning to use a novel object subjects acquired an action representation of the end location associated with using the object, as evidenced by slower responses in an action observation task, when the object was presented at an incorrect end location. By showing the importance of knowledge about end locations when learning to use a novel object, the present study suggests that end locations are a fundamental organizing feature of the human motor system.

Neural evidence for impaired action selection in right hemiparetic cerebral palsy.

Recent studies suggest that in addition to low-level motor impairments, individuals with hemiparetic cerebral palsy (HCP) are characterized by anticipatory action planning deficits as well. In the present EEG study we investigated the neural and temporal dynamics of action planning in participants with right-sided HCP (n=10) and in left-handed control subjects (n=10). An anticipatory planning task was used in which participants were required to grasp and rotate a hexagonal knob over different angles (60 degrees, 120 degrees or 180 degrees). At a behavioral level, participants with HCP were slower in their movements and often selected an inappropriate grip when grasping the object. At a neural level, individuals with HCP showed a strong reduction in the amplitude of the P2 component, likely reflecting an impaired process of action selection. In addition, a strong correlation was observed between the P2 amplitude and grasping and rotation times. The P2 component was localized to sources in the dorsal posterior cingulate cortex (dPCC), an area that is known to be involved in orienting visual body parts in space. Together these findings suggest that anticipatory planning deficits in cerebral palsy arise mainly due to an impaired process of action selection.

Neural and temporal dynamics underlying visual selection for action.

The present study investigated the selection for action hypothesis, according to which a subject's action intention to perform a movement influences the way in which visual information is being processed. Subjects were instructed in separate blocks either to grasp or to point to a three-dimensional target-object and event-related potentials were recorded relative to stimulus onset. It was found that grasping compared with pointing resulted in a stronger N1 component and a subsequent selection negativity, which were localized to the lateral occipital complex. These effects suggest that the intention to grasp influences the processing of action-relevant features in ventral stream areas already at an early stage (e.g., enhanced processing of object orientation for grasping). These findings provide new insight in the neural and temporal dynamics underlying perception-action coupling and provide neural evidence for a selection for action principle in early human visual processing.

The N400-concreteness effect reflects the retrieval of semantic information during the preparation of meaningful actions.

The present ERP study investigated when and how action semantics, i.e. the conceptual knowledge that we acquired over the course of our lives, is activated during the preparation of object-directed actions. Subjects were required to grasp one of two objects and were implicitly instructed to either perform a meaningful action (e.g. moving a cup towards the mouth) or a meaningless action with the object (e.g. moving a cup towards the eye). A larger anterior N400 was found for the preparation of meaningful compared to meaningless actions, likely reflecting the retrieval of action semantic information in case a meaningful action was required with the object. The distribution and the latency of the anterior N400-effect were strongly related to standard N400-repetition effects, thereby further corroborating the semantic nature of the effect. In sum, the present study provides new insight in the neural and temporal dynamics underlying semantics for action.

The functional role of motor activation in language processing: motor cortical oscillations support lexical-semantic retrieval.

There is increasing experimental evidence that processing action-related language results in the automatic activation of associated regions of the motor and premotor cortex. However, the functional significance of motor activation in language processing is still under debate. In the present EEG study, we set out to investigate if language-induced motor activation primarily reflects the retrieval of lexical-semantic information or post-lexical motor imagery. The processing of action verbs was found accompanied by an early activation of motor-related brain areas, as reflected by a desynchronization in the mu- and beta-frequency bands which was localized to motor and premotor areas. A stronger motor activation was observed for verbs presented in an animal context (e.g. "The deer jumped over the stream") compared to a human context (e.g. "The athlete jumped over the fence") and motor resonance was directly modulated by the cloze probability of the noun-verb pairs. The onset of the motor effects preceded classical measures of semantic integration (i.e. the N400 component) and the strength of motor activation was found inversely related to the size of the N400 effect. These findings support the hypothesis that motor activation in language processing primarily supports the retrieval and integration of lexical-semantic information.

Short-term action intentions overrule long-term semantic knowledge.

In the present study, we investigated whether the preparation of an unusual action with an object (e.g. bringing a cup towards the eye) could selectively overrule long-term semantic representations. In the first experiment it was found that unusual action intentions activated short-term semantic goal representations, rather than long-term conceptual associations. In a second experiment the reversal of long-term priming effects was replicated, while reducing the need for internal verbalization as a possible strategy to accomplish the task. Priming effects in the first two experiments were found to involve the selection of object knowledge at a semantic level, rather than reflecting a general effect of action preparation on word processing (Experiment 3). Finally, in a fourth experiment short-term priming effects were shown to extend beyond a lexical level by showing faster responses to pictures representing the short-term action goal. Together, the present findings extend the 'selection-for-action' principle previously used in visual attention to a semantic level, by showing that semantic information is selectively activated in line with the short-term goal of the actor.

You'll never crawl alone: neurophysiological evidence for experience-dependent motor resonance in infancy.

Lately, neuroscience is showing a great interest in examining the functional and neural mechanisms which support action observation and understanding. Recent studies have suggested that our motor skills crucially affect the way in which we perceive the actions generated by others, by showing stronger motor resonance for observation of actions that are established in one's motor repertoire. In the present study we extend previous findings that were based on expert motor skills in adults to the natural development of actions in infants. To investigate the effect of natural motor experience on motor resonance during action observation, 14- to 16-month-old infants' EEG was recorded during observation of action videos. Stronger mu- and beta-desynchronizations were found for observation of crawling compared to walking videos and the size of the effect was strongly related to the infant's own crawling experience. This suggests that already early in life one's own action experience is closely related to how actions of others are perceived.

Conceptual knowledge for understanding other's actions is organized primarily around action goals.

Semantic knowledge about objects entails both knowing how to grasp an object (grip-related knowledge) and what to do with an object (goal-related knowledge). Considerable evidence suggests a hierarchical organization in which specific hand-grips in action execution are most often selected to accomplish a remote action goal. The present study aimed to investigate whether a comparable hierarchical organization of semantic knowledge applies to the recognition of other's object-directed actions as well. Correctness of either the Grip (hand grip applied to the object) or the Goal (end-location at which an object was directed) were manipulated independently in two experiments. In Experiment 1, subjects were required to attend selectively to either the correctness of the grip or the goal of the observed action. Subjects were faster when attending to the goal of the action and a strong interference of goal-violations was observed when subjects attended to the grip of the action. Importantly, observation of irrelevant goal- or grip-related violations interfered with making decisions about the correctness of the relevant dimension only when the relevant dimension was correct. In contrast, in Experiment 2, when subjects attended to an action-irrelevant stimulus dimension (i.e. orientation of the object), no interference of goal- or grip-related violations was found, ruling out the possibility that interference-effects result from perceptual differences between stimuli. These findings suggest that understanding the correctness of an action selectively recruits specialized, but interacting networks, processing the correctness of goal- and grip-specific information during action observation.

Semantics in action: an electrophysiological study on the use of semantic knowledge for action.

Recent studies have supported close interactions between language and action-related processes, suggesting comparable neural mechanisms. However, relatively little is known about the semantics involved in action planning. The present study investigated the activation of semantic knowledge in meaningful actions by recording event-related potentials (ERPs). Subjects prepared meaningful or meaningless actions with objects and made a semantic categorization response before executing the action. Words presented could be either congruent or incongruent with respect to the goal of the action. Preparation of meaningful actions elicited a larger anterior N400 for words incongruent to the present action goal as compared to congruent words, while no N400 effect was found when subjects prepared meaningless actions. These findings indicate that the preparation of meaningful actions with objects is accompanied by the activation of semantic information representing the usual action goals associated with those objects.

Coordinated control of eye and hand movements in dynamic reaching.

In the present study, we integrated two recent, at first sight contradictory findings regarding the question whether saccadic eye movements can be generated to a newly presented target during an ongoing hand movement. Saccades were measured during so-called adaptive and sustained pointing conditions. In the adapted pointing condition, subjects had to direct both their gaze and arm movements to a displaced target location. The results showed that the eyes could fixate the new target during pointing. In addition, a temporal coupling of these corrective saccades was found with changes in arm movement trajectories when reaching to the new target. In the sustained pointing condition, however, the same subjects had to point to the initial target, while trying to deviate their gaze to a new target that appeared during pointing. It was found that the eyes could not fixate the new target before the hand reached the initial target location. Together, the results indicate that ocular gaze is always forced to follow the target intended by a manual arm movement. A neural mechanism is proposed that couples ocular gaze to the target of an arm movement. Specifically, the mechanism includes a reach neuron layer besides the well-known saccadic layer in the primate superior colliculus. Such a tight, sub-cortical coupling of ocular gaze to the target of a reaching movement can explain the contrasting behavior of the eyes in dependency of whether the eye and hand share the same target position or attempt to move to different locations.