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G Rizzolatti

Publications and source records attributed to G Rizzolatti.

At least 19 recordsLinked to original sources

Listening to action-related sentences modulates the activity of the motor system: a combined TMS and behavioral study.

Transcranial magnetic stimulation (TMS) and a behavioral paradigm were used to assess whether listening to action-related sentences modulates the activity of the motor system. By means of single-pulse TMS, either the hand or the foot/leg motor area in the left hemisphere was stimulated in distinct experimental sessions, while participants were listening to sentences expressing hand and foot actions. Listening to abstract content sentences served as a control. Motor evoked potentials (MEPs) were recorded from hand and foot muscles. Results showed that MEPs recorded from hand muscles were specifically modulated by listening to hand-action-related sentences, as were MEPs recorded from foot muscles by listening to foot-action-related sentences. This modulation consisted of an amplitude decrease of the recorded MEPs. In the behavioral task, participants had to respond with the hand or the foot while listening to actions expressing hand and foot actions, as compared to abstract sentences. Coherently with the results obtained with TMS, when the response was given with the hand, reaction times were slower during listening to hand-action-related sentences, while when the response was given with the foot, reaction times were slower during listening to foot-action-related sentences. The present data show that processing verbally presented actions activates different sectors of the motor system, depending on the effector used in the listened-to action.

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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 one's 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 imitator's 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.

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The cortical motor system.

The cortical motor system of primates is formed by a mosaic of anatomically and functionally distinct areas. These areas are not only involved in motor functions, but also play a role in functions formerly attributed to higher order associative cortical areas. In the present review, we discuss three types of higher functions carried out by the motor cortical areas: sensory-motor transformations, action understanding, and decision processing regarding action execution. We submit that generating internal representations of actions is central to cortical motor function. External contingencies and motivational factors determine then whether these action representations are transformed into actual actions.

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I know what you are doing. a neurophysiological study.

In the ventral premotor cortex of the macaque monkey, there are neurons that discharge both during the execution of hand actions and during the observation of the same actions made by others (mirror neurons). In the present study, we show that a subset of mirror neurons becomes active during action presentation and also when the final part of the action, crucial in triggering the response in full vision, is hidden and can therefore only be inferred. This implies that the motor representation of an action performed by others can be internally generated in the observer's premotor cortex, even when a visual description of the action is lacking. The present findings support the hypothesis that mirror neuron activation could be at the basis of action recognition.

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Cortical mechanism for the visual guidance of hand grasping movements in the monkey: A reversible inactivation study.

Picking up an object requires two basic motor operations: reaching and grasping. Neurophysiological studies in monkeys have suggested that the visuomotor transformations necessary for these two operations are carried out by separate parietofrontal circuits and that, for grasping, a key role is played by a specific sector of the ventral premotor cortex: area F5. The aim of the present study was to test the validity of this hypothesis by reversibly inactivating area F5 in monkeys trained to grasp objects of different shape, size and orientation. In separate sessions, the hand field of the primary motor cortex (area F1 or area 4) was also reversibly inactivated. The results showed that after inactivation of area F5 buried in the bank of the arcuate sulcus (the F5 sector where visuomotor neurones responding to object presentation are located), the hand shaping preceding grasping was markedly impaired and the hand posture was not appropriate for the object size and shape. The monkeys were eventually able to grasp the objects, but only after a series of corrections made under tactile control. With small inactivations the deficits concerned the contralesional hand, with larger inactivations the ipsilateral hand as well. In addition, there were signs of peripersonal neglect in the hemispace contralateral to the inactivation site. Following inactivation of area F5 lying on the cortical convexity (the F5 sector where visuomotor neurones responding to action observation, 'mirror neurones', are found) only a motor slowing was observed, the hand shaping being preserved. The inactivation of the hand field of area F1 produced a severe paralysis of contralateral finger movements with hypotonia. The results of this study indicate the crucial role of the ventral premotor cortex in visuomotor transformations for grasping movements. More generally, they provide strong support for the notion that distal and proximal movement organization relies upon distinct cortical circuits. Clinical data on distal movement deficits in humans are re-examined in the light of the present findings.

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Action observation activates premotor and parietal areas in a somatotopic manner: an fMRI study.

Functional magnetic resonance imaging (fMRI) was used to localize brain areas that were active during the observation of actions made by another individual. Object- and non-object-related actions made with different effectors (mouth, hand and foot) were presented. Observation of both object- and non-object-related actions determined a somatotopically organized activation of premotor cortex. The somatotopic pattern was similar to that of the classical motor cortex homunculus. During the observation of object-related actions, an activation, also somatotopically organized, was additionally found in the posterior parietal lobe. Thus, when individuals observe an action, an internal replica of that action is automatically generated in their premotor cortex. In the case of object-related actions, a further object-related analysis is performed in the parietal lobe, as if the subjects were indeed using those objects. These results bring the previous concept of an action observation/execution matching system (mirror system) into a broader perspective: this system is not restricted to the ventral premotor cortex, but involves several somatotopically organized motor circuits.

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Visuomotor neurons: ambiguity of the discharge or 'motor' perception?

The cortical motor system has been classically considered as the unitary, output stage of the brain processing of sensory information. According to this idea, the motor cortex - the acting brain - receives the result of the perceptual processing (visual, acoustical, tactile, etc.) elaborated by the 'associative cortex'. During the last two decades this perspective has been challenged by a series of anatomical, hodological, and neurophysiological data. This converging evidence delineates a dramatically changed picture. Far from being unitary, the cortical motor system appears to be constituted by a constellation of distinct areas, each of those endowed with specific functional properties and linked by reciprocal connections with distinct sectors of the parietal cortex. Furthermore, several 'motor' neurons in addition to their motor discharge, are also activated by somatosensory and visual stimulation (somatomotor and visuomotor neurons). In the present paper we will discuss the functional properties of those sensorimotor neurons located in the ventral part of the monkey premotor cortex. On the basis of electrophysiological data, we will propose that the apparent parodox stemming from the coexistence within the same neuron of motor and sensory properties can be solved by postulating that the motor system not only executes actions but also internally represents them in terms of 'motor ideas'. These motor ideas may provide the neurobiological basis for space representation, understanding of actions made by others and, possibly, semantic categorization of objects.

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Cortical mechanisms of human imitation.

How does imitation occur? How can the motor plans necessary for imitating an action derive from the observation of that action? Imitation may be based on a mechanism directly matching the observed action onto an internal motor representation of that action ("direct matching hypothesis"). To test this hypothesis, normal human participants were asked to observe and imitate a finger movement and to perform the same movement after spatial or symbolic cues. Brain activity was measured with functional magnetic resonance imaging. If the direct matching hypothesis is correct, there should be areas that become active during finger movement, regardless of how it is evoked, and their activation should increase when the same movement is elicited by the observation of an identical movement made by another individual. Two areas with these properties were found in the left inferior frontal cortex (opercular region) and the rostral-most region of the right superior parietal lobule.

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A parieto-premotor network for object manipulation: evidence from neuroimaging.

Functional magnetic resonance imaging (fMRI) was used to assess cerebral activation during manipulation of various complex meaningless objects as compared to manipulation of a single simple object (a sphere). Significant activation was found bilaterally in the ventral premotor cortex (Brodmann's area 44), in the cortex lining the anterior part of the intraparietal sulcus (most probably corresponding to monkey anterior intraparietal area, AIP), in the superior parietal lobule and in the opercular parietal cortex including the secondary somatosensory area (SII). We suggest that the cortex lining the anterior part of the intraparietal sulcus and area 44 are functionally connected and mediate object manipulation in humans.

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Action for perception: a motor-visual attentional effect.

Five experiments investigated whether preparation of a grasping movement affects detection and discrimination of visual stimuli. Normal human participants were required to prepare to grasp a bar and then to grasp it as fast as possible on presentation of a visual stimulus. On the basis of the degree of sharing of their intrinsic properties with those of the to-be-grasped bar, visual stimuli were categorized as "congruent" or "incongruent." Results showed that grasping reaction times to congruent visual stimuli were faster than reaction times to incongruent ones. These data indicate that preparation to act on an object produces faster processing of stimuli congruent with that object. The same facilitation was present also when, after the preparation of hand grasping, participants were suddenly instructed to inhibit the prepared grasping movement and to respond with a different motor effector. The authors suggest that these findings could represent an extension of the premotor theory of attention, from orienting of attention to spatial locations to orienting of attention to graspable objects.

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A fronto-parietal circuit for object manipulation in man: evidence from an fMRI-study.

Functional magnetic resonance imaging (fMRI) was used to localize brain areas active during manipulation of complex objects. In one experiment subjects were required to manipulate complex objects for exploring their macrogeometric features as compared to manipulation of a simple smooth object (a sphere). In a second experiment subjects were asked to manipulate complex objects and to silently name them upon recognition as compared to manipulation of complex not recognizable objects without covert naming. Manipulation of complex objects resulted in an activation of ventral premotor cortex [Brodmann's area (BA) 44], of a region in the intraparietal sulcus (most probably corresponding to the anterior intraparietal area in the monkey), of area SII and of a sector of the superior parietal lobule. When the objects were covertly named additional activations were found in the opercular part of BA 44 and in the pars triangularis of the inferior frontal gyrus (BA 45). We suggest that a fronto-parietal circuit for manipulation of objects exists in humans and involves basically the same areas as in the monkey. It is proposed that area SII analyses the intrinsic object characteristics whilst the superior parietal lobule is related to kinaesthesia.

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Resonance behaviors and mirror neurons.

This article is subdivided into two parts. In the first part we review the properties of a particular class of premotor neurons, the "mirror" neurons. With this term we define neurons that discharge both when the monkey makes a particular action and when it observes another individual (monkey or human) making a similar action. The second part is an attempt to give a neurophysiological account of the mechanisms underlying behaviors where an individual reproduces, overtly or internally, movements or actions made by another individual. We will refer to these behaviors as "resonance behaviors". We distinguish two types of resonance behavior. The first type is characterized by imitation, immediate or with delay, of movements made by other individuals. Examples of resonance behavior of this type are the "imitative" behaviors observed in birds, young infants and patients with frontal lesions. The second type of resonance behavior is characterized by the occurrence, at the observation of an action, of a neural pattern, which, when internally generated, determines the making of the observed action. In this type of resonance behavior the observed action is, typically, not repeated (overtly). We argue that resonance behavior of the second type is at the basis of the understanding of actions made by others. At the end of the article we review evidence of mirror mechanisms in humans and discuss their anatomical localizations.

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Activation of human primary motor cortex during action observation: a neuromagnetic study.

The monkey premotor cortex contains neurons that discharge during action execution and during observation of actions made by others. Transcranial magnetic stimulation experiments suggest that a similar observation/execution matching system also is present in humans. We recorded neuromagnetic oscillatory activity of the human precentral cortex from 10 healthy volunteers while (i) they had no task to perform, (ii) they were manipulating a small object, and (iii) they were observing another individual performing the same task. The left and right median nerves were stimulated alternately (interstimulus interval, 1.5 s) at intensities exceeding motor threshold, and the poststimulus rebound of the rolandic 15- to 25-Hz activity was quantified. In agreement with previous studies, the rebound was strongly suppressed bilaterally during object manipulation. Most interestingly, the rebound also was significantly diminished during action observation (31-46% of the suppression during object manipulation). Control experiments, in which subjects were instructed to observe stationary or moving stimuli, confirmed the specificity of the suppression effect. Because the recorded 15- to 25-Hz activity is known to originate mainly in the precentral motor cortex, we concluded that the human primary motor cortex is activated during observation as well as execution of motor tasks. These findings have implications for a better understanding of the machinery underlying action recognition in humans.

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Grasping objects and grasping action meanings: the dual role of monkey rostroventral premotor cortex (area F5).

Monkey area F5 consists of two main histochemical sectors, one buried inside the arcuate sulcus, the other located on the cortical convexity. Neurons of both sectors discharge during hand movements. Many of them also fire in response to the presentation of visual stimuli. However, the visual stimuli effective for triggering the neurons in each sector are markedly different. Neurons located in the bank of the arcuate sulcus respond to the observation of 3D objects, provided that object size and shape is congruent with the prehension type coded by the neuron ('canonical' F5 neurons). Neurons of the convexity discharge when the monkey observes hand actions performed by another individual, provided that they are similar to the motor action coded by the neuron ('mirror' neurons). What do the canonical F5 neurons and the surprising mirror neurons have in common? The interpretation we propose is that these two categories of F5 neurons both generate an internal copy of a potential hand action. In the case of 'canonical' neurons, this copy gives a description of how to grasp an object; in the case of mirror neurons it gives a description of an action made by another person. Because the individuals know the consequences of their actions, we propose that the internal motor copies of the observed actions represent the neural basis for understanding the meaning of actions made by others.

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The organization of the cortical motor system: new concepts.

A series of recent anatomical and functional data has radically changed our view on the organization of the motor cortex in primates. In the present article we present this view and discuss its fundamental principles. The basic principles are the following: (a) the motor cortex, defined as the agranular frontal cortex, is formed by a mosaic of separate areas, each of which contains an independent body movement representation, (b) each motor area plays a specific role in motor control, based on the specificity of its cortical afferents and descending projections, (c) in analogy to the motor cortex, the posterior parietal cortex is formed by a multiplicity of areas, each of which is involved in the analysis of particular aspects of sensory information. There are no such things as multipurpose areas for space or body schema and (d) the parieto-frontal connections form a series of segregated anatomical circuits devoted to specific sensorimotor transformations. These circuits transform sensory information into action. They represent the basic functional units of the motor system. Although these conclusions mostly derive from monkey experiments, anatomical and brain-imaging evidence suggest that the organization of human motor cortex is based on the same principles. Possible homologies between the motor cortices of humans and non-human primates are discussed.

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Language within our grasp.

In monkeys, the rostral part of ventral premotor cortex (area F5) contains neurons that discharge, both when the monkey grasps or manipulates objects and when it observes the experimenter making similar actions. These neurons (mirror neurons) appear to represent a system that matches observed events to similar, internally generated actions, and in this way forms a link between the observer and the actor. Transcranial magnetic stimulation and positron emission tomography (PET) experiments suggest that a mirror system for gesture recognition also exists in humans and includes Broca's area. We propose here that such an observation/execution matching system provides a necessary bridge from'doing' to'communicating',as the link between actor and observer becomes a link between the sender and the receiver of each message.

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