Mirror neurons are a specific type of visuomotor neuron that discharge both when a monkey executes a motor act and when it observes a similar motor act performed by another individual. In this ...article, we review first the basic properties of these neurons. We then describe visual features recently investigated which indicate that, besides encoding the goal of motor acts, mirror neurons are modulated by location in space of the observed motor acts, by the perspective from which the others’ motor acts are seen, and by the value associated with the object on which others’ motor acts are performed. In the last part of this article, we discuss the role of the mirror mechanism in planning actions and in understanding the intention underlying the others’ motor acts. We also review some human studies suggesting that motor intention in humans may rely, as in the monkey, on the mirror mechanism.
Studies on action observation mostly described the activation of a network of cortical areas, while less investigation focused specifically on the activation and role of subcortical nodes. In the ...present fMRI study, we investigated the recruitment of cerebellum and basal ganglia during the execution and observation of object manipulation performed with the right hand. The observation conditions consisted in: (a) observation of manipulative actions; (b) observation of sequences of random finger movements. In the execution conditions, participants had to perform the same actions or movements as in (a) and (b), respectively. The results of conjunction analysis showed significant shared activations during both observation and execution of manipulation in several subcortical structures, including: (1) cerebellar lobules V, VI, crus I, VIIIa and VIIIb (bilaterally); (2) globus pallidus, bilaterally, and left subthalamic nucleus; (3) red nucleus (bilaterally) and left thalamus. These findings support the hypothesis that the action observation/execution network also involves subcortical structures, such as cerebellum and basal ganglia, forming an integrated network. This suggests possible mechanisms, involving these subcortical structures, underlying learning of new motor skills, through action observation and imitation.
The observation of actions performed by others is believed to activate the Action Observation Network (AON). Previous evidence suggests that subjects with a specific motor skill show increased ...activation of the AON during observation of the same skill. The question arises regarding which modulation of the AON occurs during observation of novel complex manipulative actions that are beyond the personal motor repertoire. To address this issue, we carried out a functional MRI study in which healthy volunteers without specific hand motor skills observed videos displaying hand-object manipulation executed by an expert with high manual dexterity, by an actor with intermediate ability or by a naïve subject. The results showed that the observation of actions performed by a naïve model produced stronger activation in a dorso-medial parieto-premotor circuit including the superior parietal lobule and dorsal premotor cortex, compared to observation of an expert actor. Functional connectivity analysis comparing the observation of the naïve model with that of the expert model, revealed increased connectivity between dorsal areas of the AON. This suggests a possible distinction between ventral and dorsal brain circuits involved in the processing of different aspects of action perception, such as kinematics and final action goal.
The macaque ventral premotor area F5 hosts two types of visuomotor grasping neurons: "canonical" neurons, which respond to visually presented objects and underlie visuomotor transformation for ...grasping, and "mirror" neurons, which respond during the observation of others' action, likely playing a role in action understanding. Some previous evidence suggested that canonical and mirror neurons could be anatomically segregated in different sectors of area F5. Here we investigated the functional properties of single neurons in the hand field of area F5 using various tasks similar to those originally designed to investigate visual responses to objects and actions. By using linear multielectrode probes, we were able to simultaneously record different types of neurons and to precisely localize their cortical depth. We recorded 464 neurons, of which 243 showed visuomotor properties. Canonical and mirror neurons were often present in the same cortical sites; and, most interestingly, a set of neurons showed both canonical and mirror properties, discharging to object presentation as well as during the observation of experimenter's goal-directed acts (canonical-mirror neurons). Typically, visual responses to objects were constrained to the monkey peripersonal space, whereas action observation responses were less space-selective. Control experiments showed that space-constrained coding of objects mostly relies on an operational (action possibility) rather than metric (absolute distance) reference frame. Interestingly, canonical-mirror neurons appear to code object as target for both one's own and other's action, suggesting that they could play a role in predictive representation of others' impending actions.
Starting from the proposed role of the mirror neuron system in the recognition of the intention underlying the actions of others, an experimental paradigm was implemented to test the role of sailing ...motor expertise in predicting the outcome of a competitor's action. It was hypothesized that subjects with experience in sailing would correctly interpret the maneuver performed due to the activation of domain specific motor representations of the same movements and that subjects who practiced a sport different from sailing would perform worse because of the activation of irrelevant motor patterns. For doing so, a series of video clips, in which a professional sailor performed a tack or a feint, have been manipulated so that the video clips would stop at the moment of the dunkin, namely, when the boat acquires speed to tack or continue straight ahead. The task consisted in predicting whether the action following the dunkin was an actual tack or a feint. The performance of 87 subjects, divided into three subgroups (sailors, tennis players, sedentary), was evaluated in terms of accuracy in identifying the sailor's intentions and correlated to age, gender, manual dominance, education, job, hours spent weekly playing videogames, and experience in playing sports. Results showed that the percentage of correct identifications of the intention to do a tack or feint was the highest in the group of sailors and the lowest in tennis players. An inverse relation between tennis experience and ability in recognizing the sailor's intention was found in the group of tennis players. Gender, age, manual dominance, education, job, and experience with videogames were not found to be correlated with performance. Findings support the possible implication of the mirror neuron system in maneuver detection in sailing and may be a starting point for the development of psychological training in this sport.
Information about objects around us is essential for planning actions and for predicting those of others. Here, we studied pre-supplementary motor area F6 neurons with a task in which monkeys viewed ...and grasped (or refrained from grasping) objects, and then observed a human doing the same task. We found “action-related neurons” encoding selectively monkey’s own action self-type (ST), another agent’s action other-type (OT), or both self- and other-type (SOT). Interestingly, we found “object-related neurons” exhibiting the same type of selectivity before action onset: Indeed, distinct sets of neurons discharged when visually presented objects were targeted by the monkey’s own action (ST), another agent’s action (OT), or both (SOT). Notably, object-related neurons appear to signal self and other’s intention to grasp and the most likely grip type that will be performed, whereas action-related neurons encode a general goal attainment signal devoid of any specificity for the observed grip type. Time-resolved cross-modal population decoding revealed that F6 neurons first integrate information about object and context to generate an agent-shared signal specifying whether and how the object will be grasped, which progressively turns into a broader agent-based goal attainment signal during action unfolding. Importantly, shared representation of objects critically depends upon their location in the observer’s peripersonal space, suggesting an “object-mirroring” mechanism through which observers could accurately predict others’ impending action by recruiting the same motor representation they would activate if they were to act upon the same object in the same context.
The ventral part of lateral prefrontal cortex (VLPF) of the monkey receives strong visual input, mainly from inferotemporal cortex. It has been shown that VLPF neurons can show visual responses ...during paradigms requiring to associate arbitrary visual cues to behavioral reactions. Further studies showed that there are also VLPF neurons responding to the presentation of specific visual stimuli, such as objects and faces. However, it is largely unknown whether VLPF neurons respond and differentiate between stimuli belonging to different categories, also in absence of a specific requirement to actively categorize or to exploit these stimuli for choosing a given behavior. The first aim of the present study is to evaluate and map the responses of neurons of a large sector of VLPF to a wide set of visual stimuli when monkeys simply observe them. Recent studies showed that visual responses to objects are also present in VLPF neurons coding action execution, when they are the target of the action. Thus, the second aim of the present study is to compare the visual responses of VLPF neurons when the same objects are simply observed or when they become the target of a grasping action. Our results indicate that: (1) part of VLPF visually responsive neurons respond specifically to one stimulus or to a small set of stimuli, but there is no indication of a "passive" categorical coding; (2) VLPF neuronal visual responses to objects are often modulated by the task conditions in which the object is observed, with the strongest response when the object is target of an action. These data indicate that VLPF performs an early passive description of several types of visual stimuli, that can then be used for organizing and planning behavior. This could explain the modulation of visual response both in associative learning and in natural behavior.
A fundamental capacity of social animals consists in the predictive representation of upcoming events in the outside world, such as the actions of others. Here, we tested the activity of ventral ...premotor area F5 mirror neurons (MNs) while monkeys observed an experimenter performing (Action condition) or withholding (Inaction condition) a grasping action, which could be predicted on the basis of previously presented auditory instructions. Many of the recorded MNs discharged only during action observation (Action MNs), but one-third also encoded the experimenter's withheld action (Inaction MNs). Interestingly, while most of Action MNs exhibited reactive discharge during action observation, becoming active after the go signal, the majority of Inaction MNs showed predictive discharge. MN population activity as a whole displayed an overall predictive activation pattern, becoming active, on average, 340 ms before the go signal. Furthermore, MNs became active earlier when the observed action was performed in the monkeys' extrapersonal rather than peripersonal space, suggesting that context-based neural prediction of others' actions plays different roles depending on the monkeys' ability to interact with the observed agent.
What are the neural bases of action understanding? Although this capacity could merely involve visual analysis of the action, it has been argued that we actually map this visual information onto its ...motor representation in our nervous system. Here we discuss evidence for the existence of a system, the 'mirror system', that seems to serve this mapping function in primates and humans, and explore its implications for the understanding and imitation of action.
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DOBA, IJS, IZUM, KILJ, NUK, PILJ, PNG, SAZU, UILJ, UKNU, UL, UM, UPUK
Inferior parietal lobule (IPL) neurons were studied when monkeys performed motor acts embedded in different actions and when they observed similar acts done by an experimenter. Most motor IPL neurons ...coding a specific act (e.g., grasping) showed markedly different activations when this act was part of different actions (e.g., for eating or for placing). Many motor IPL neurons also discharged during the observation of acts done by others. Most responded differentially when the same observed act was embedded in a specific action. These neurons fired during the observation of an act, before the beginning of the subsequent acts specifying the action. Thus, these neurons not only code the observed motor act but also allow the observer to understand the agent's intentions.
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