Harold Bekkering

Rational imitation in preverbal infants

Here we show that if an adult demonstrates a new way to execute a task to a group of infants aged 14 months, the children will use this action to achieve the same goal only if they consider it to be the most rational alternative. Our results indicate that imitation of goal-directed action by preverbal infants is a selective, interpretative process, rather than a simple re-enactment of the means used by a demonstrator, as was previously thought.

Movement observation affects movement execution in a simple response task

The present study was designed to examine the hypothesis that stimulus-response arrangements with high ideomotor compatibility lead to substantial compatibility effects even in simple response tasks. In Experiment 1, participants executed pre-instructed finger movements in response to compatible and incompatible finger movements. A pronounced reaction time advantage was found for compatible as compared to incompatible trials. Experiment 2 revealed a much smaller compatibility effect for less ideomotor-compatible object movements compared to finger movements. Experiment 3 presented normal stimuli (hand upright) and flipped stimuli (hand upside-down). Two components were found to contribute to the compatibility effect, a dynamic spatial compatibility component (related to movement directions) and an ideomotor component (related to movement types). The implications of these results for theories about stimulus-response compatibility (SRC) as well as for theories about imitation are discussed.

Compatibility between observed and executed finger movements: Comparing symbolic, spatial, and imitative cues

Intuitively, one can assume that imitating a movement is an easier task than responding to a symbolic stimulus like a verbal instruction. Support for this suggestion can be found in neuropsychological research as well as in research on stimulus-response compatibility. However controlled experimental evidence for this assumption is still lacking. We used a stimulus-response compatibility paradigm to test the assumption. In a series of experiments, it was tested whether observed finger movements have a stronger influence on finger movement execution than a symbolic or spatial cue. In the first experiment, we compared symbolic cues with observed finger movements using an interference paradigm. Observing finger movements strongly influenced movement execution, irrespective of whether the finger movement was the relevant or the irrelevant stimulus dimension. In the second experiment, effects of observed finger movements and spatial finger cues were compared. The observed finger movement dominated the spatial finger cue. A reduction in the similarity of observed and executed action in the third experiment led to a decrease of the influence of observed finger movement, which demonstrates the crucial role of the imitative relation of observed and executed action for the described effects. The results are discussed in relation to recent models of stimulus-response compatibility. Neurocognitive support for the strong relationship between movement observation and movement execution is reported.

Imitation of Gestures in Children is Goal-directed:

The view that the motor program activated during imitation is organized by goals was investigated by asking pre-school children to imitate a set of hand gestures of varying complexity that were made by an experimenter sitting in front of them. In Experiments 1 and 3, children reached for the correct object (one of their own ears or one of two dots on a table) but preferred to use the ipsilateral hand. This ipsilateral preference was not observed when hand movements were made to only one ear (Experiment 2), or when movements were diercted at space rather than physical objects (Experiment 3). The results are consistent with the notion that imitation is guided by goals and provide insights about how these goals are organized.

Reafferent copies of imitated actions in the right superior temporal cortex

Imitation is a complex phenomenon, the neural mechanisms of which are still largely unknown. When individuals imitate an action that already is present in their motor repertoire, a mechanism matching the observed action onto an internal motor representation of that action should suffice for the purpose. When one has to copy a new action, however, or to adjust an action present in ones motor repertoire to a different observed action, an additional mechanism is needed that allows the observer to compare the action made by another individual with the sensory consequences of the same action made by himself. Previous experiments have shown that a mechanism that directly matches observed actions on their motor counterparts exists in the premotor cortex of monkeys and humans. Here we report the results of functional magnetic resonance experiments, suggesting that in the superior temporal sulcus, a higher order visual region, there is a sector that becomes active both during hand action observation and during imitation even in the absence of direct vision of the imitators hand. The motor-related activity is greater during imitation than during control motor tasks. This newly identified region has all the requisites for being the region at which the observed actions, and the reafferent motor-related copies of actions made by the imitator, interact.

Modulation of activity in medial frontal and motor cortices during error observation

We used measures of the human event-related brain potential (ERP) to investigate the neural mechanisms underlying error processing during action observation. Participants took part in two conditions, a task execution condition and a task observation condition. We found that activity in both the medial frontal cortex and the motor cortices, as measured via the error-related negativity and the lateralized readiness potential, respectively, was modulated by the correctness of observed behavior. These data suggest that similar neural mechanisms are involved in monitoring ones own actions and the actions of others.

The mirror neuron system is more active during complementary compared with imitative action

We assessed the role of the human mirror neuron system (MNS) in complementary actions using functional magnetic resonance imaging while participants prepared to execute imitative or complementary actions. The BOLD signal in the right inferior frontal gyrus and bilateral inferior parietal lobes was greater during preparation of complementary than during imitative actions, suggesting that the MNS may be essential in dynamically coupling action observation to action execution.

Visual Search Is Modulated by Action Intentions

The influence of action intentions on visual selection processes was investigated in a visual search paradigm. A predefined target object with a certain orientation and color was presented among distractors, and subjects had to either look and point at the target or look at and grasp the target. Target selection processes prior to the first saccadic eye movement were modulated by the different action intentions. Specifically, fewer saccades to objects with the wrong orientation were made in the grasping condition than in the pointing condition, whereas the number of saccades to an object with the wrong color was the same in the two conditions. Saccadic latencies were similar under the different task conditions, so the results cannot be explained by a speed-accuracy trade-off. The results suggest that a specific action intention, such as grasping, can enhance visual processing of action-relevant features, such as orientation. Together, the findings support the view that visual attention can be best understood as a selection-for-action mechanism.

Parieto-Frontal Connectivity during Visually Guided Grasping

Grasping an object requires processing visuospatial information about the extrinsic features (spatial location) and intrinsic features (size, shape, orientation) of the object. Accordingly, manual prehension has been subdivided into a reach component, guiding the hand toward the object on the basis of its extrinsic features, and a grasp component, preshaping the fingers around the center of mass of the object on the basis of its intrinsic features. In neural terms, this distinction has been linked to a dedicated dorsomedial “reaching” circuit and a dorsolateral “grasping” circuit that process extrinsic and intrinsic features, linking occipital areas via parietal regions with the dorsal and ventral premotor cortex, respectively. We have tested an alternative possibility, namely that the relative contribution of the two circuits is related to the degree of on-line control required by the prehension movement. We used dynamic causal modeling of functional magnetic resonance imaging time series to assess how parieto-frontal connectivity is modulated by planning and executing prehension movements toward objects of different size and width. This experimental manipulation evoked different movements, with different planning and execution phases for the different objects. Crucially, grasping large objects increased inter-regional couplings within the dorsomedial circuit, whereas grasping small objects increased the effective connectivity of a mainly dorsolateral circuit, with a degree of overlap between these circuits. These results argue against the presence of dedicated cerebral circuits for reaching and grasping, suggesting that the contributions of the dorsolateral and the dorsomedial circuits are a function of the degree of on-line control required by the movement.

Children's coding of human action: Cognitive factors influencing imitation in 3-year-olds

We used imitation as a tool for investigating how young children code action. The study was designed to examine the errors children make in re-enacting manual gestures they see. Thirty-two 3-year-old children served as subjects. Each child was shown 24 gestures, generated by systematically crossing four factors: visual monitoring, spatial endpoint, movement path, and number of hands. The results showed no difference as a function of whether the children could visually monitor their own responses. Interestingly, children made significantly more errors when the adults action terminated on a body part than they did when the same movement terminated near the body part. There were also significantly more errors when the demonstrated act involved crossing midline than when it did not, and more errors when it involved one hand rather than two hands. Our hypothesis is that human acts are coded in terms of goals. The goals are hierarchically organized, and because young children have difficulty simultaneously integrating multiple goals into one act they often re-enact the goals that are ranked higher, which leads to the errors observed. We argue that imitation is an active reconstruction of perceived events and taps cognitive processing. We suggest that the goal-based imitation in 3-year-olds is a natural developmental outgrowth of the perceptual-motor mapping and goal-directed coding of human acts found in infancy.