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Biomedical subjects

L Fogassi

Publications and source records attributed to L Fogassi.

At least 19 recordsLinked to original sources

The observation and hearing of eating actions activates motor programs related to eating in macaque monkeys.

The observation of actions can lead, in some cases, to the repetition of those same actions. In other words, motor programs similar to those observed can be recruited. Since this phenomenon is expressed when in the presence of another individual, it has been named social facilitation. In the present study we investigated whether the observation and/or hearing of eating actions facilitate eating behaviors in observing/listening pig-tailed macaques. In experiment 1, the observation of an eating room mate significantly enhanced eating behavior in the observer. Similar results were obtained (experiment 2) in response to the sound of eating actions but not to control sounds (experiment 3). We propose that eating facilitation triggered by observation or listening of eating actions can rely on the mirror neuron system of ventral premotor cortex that provides a matching between the observed/listened action and the executed action. This matching system can subsequently trigger the motor programs necessary for repeating the observed/heard actions.

Acoustic Stimulation↗

Audiovisual mirror neurons and action recognition.

Many object-related actions can be recognized both by their sound and by their vision. Here we describe a population of neurons in the ventral premotor cortex of the monkey that discharge both when the animal performs a specific action and when it hears or sees the same action performed by another individual. These 'audiovisual mirror neurons' therefore represent actions independently of whether these actions are performed, heard or seen. The magnitude of auditory and visual responses did not differ significantly in half the neurons. A neurometric analysis revealed that based on the response of these neurons, two actions could be discriminated with 97% accuracy.

Acoustic Stimulation↗

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.

Animals↗

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.

Animals↗

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.

Adult↗

The ability to follow eye gaze and its emergence during development in macaque monkeys.

The ability of monkeys to follow the gaze of other individuals is a matter of debate in many behavioral studies. Physiological studies have shown that in monkeys, as in humans, there are neural correlates of eye direction detection. There is little evidence at the behavioral level, however, of the presence and development of such abilities in monkeys. The aim of the present study was to assess in juveniles and adult pig-tailed macaques (Macaca nemestrina) the capacity to use eye cues only to follow the gaze of an experimenter. Biological stimuli (head, eye, and trunk movements) were presented by an experimenter to 2 adult monkeys with their heads restrained (Experiment 1) and to 11 monkeys of different ages, free to move in their home cages (Experiment 2). A nonbiological stimulus served as a control. Results showed that macaques can follow the gaze of the experimenter by using head/eye and eye cues alone. Trunk movements and nonbiological stimuli did not significantly elicit similar reactions. Juvenile monkeys were not able to orient their attention on the basis of eye cues alone. In general, gaze following was more frequent in adults than in juveniles. Like in humans, however, such abilities in macaques dramatically improve with age suggesting that the transition to adulthood is a crucial period in the development of gaze-following behavior.

Animals↗

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.

Animals↗

Visual responses in the dorsal premotor area F2 of the macaque monkey.

This study aimed to determine the presence of neurons responding to visual stimuli in area F2 of the dorsal premotor cortex of the macaque monkey. In order to delimit the sector in which visually responsive neurons are located, the somatotopic organization of area F2 was studied with intracortical microstimulation and single neuron recording. The results showed that: (1) in area F2 there is a significant percentage of visually responsive neurons (15.9% of all recorded neurons); (2) area F2 is excitable with a low-threshold current (average 28.1 microA) and has a somatotopic representation of the whole body, except the face; and (3) most visually driven neurons (n=130 out of 169) are concentrated within the rostrolateral sector of the forelimb representation of area F2, thus providing for the first time functional support for the neuroanatomical evidence that the visual input to area F2 is mostly restricted to this sector.

Animals↗

Corticospinal excitability is specifically modulated by motor imagery: a magnetic stimulation study.

Transcranial magnetic stimulation (TMS) was used to investigate whether the excitability of the corticospinal system is selectively affected by motor imagery. To this purpose, we performed two experiments. In the first one we recorded motor evoked potentials from right hand and arm muscles during mental simulation of flexion/extension movements of both distal and proximal joints. In the second experiment we applied magnetic stimulation to the right and the left motor cortex of subjects while they were imagining opening or closing their right or their left hand. Motor evoked potentials (MEPs) were recorded from a hand muscle contralateral to the stimulated cortex. The results demonstrated that the excitability pattern during motor imagery dynamically mimics that occurring during movement execution. In addition, while magnetic stimulation of the left motor cortex revealed increased corticospinal excitability when subjects imagined ipsilateral as well as contralateral hand movements, the stimulation of the right motor cortex revealed a facilitatory effect induced by imagery of contralateral hand movements only. In conclusion, motor imagery is a high level process, which, however, manifests itself in the activation of those same cortical circuits that are normally involved in movement execution.

Adult↗

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.

Animals↗

Parietal cortex: from sight to action.

Recent findings have altered radically our thinking about the functional role of the parietal cortex. According to this view, the parietal lobe consists of a multiplicity of areas with specific connections to the frontal lobe. These areas, together with the frontal areas to which they are connected, mediate distinct sensorimotor transformations related to the control of hand, arm, eye or head movements. Space perception is not unitary, but derives from the joint activity of the fronto-parietal circuits that control actions requiring space computation.

Animals↗

Object representation in the ventral premotor cortex (area F5) of the monkey.

Visual and motor properties of single neurons of monkey ventral premotor cortex (area F5) were studied in a behavioral paradigm consisting of four conditions: object grasping in light, object grasping in dark, object fixation, and fixation of a spot of light. The employed objects were six different three-dimensional (3-D) geometric solids. Two main types of neurons were distinguished: motor neurons (n = 25) and visuomotor neurons (n = 24). Motor neurons discharged in association with grasping movements. Most of them (n = 17) discharged selectively during a particular type of grip. Different objects, if grasped in similar way, determined similar neuronal motor responses. Visuomotor neurons also discharged during active movements, but, in addition, they fired also in response to the presentation of 3-D objects. The majority of visuomotor neurons (n = 16) showed selectivity for one or few objects. The response was present both in object grasping in light and in object fixation conditions. Visuomotor neurons that selectively discharged to the presentation of a given object discharged also selectively during grasping of that object. In conclusion, object shape is coded in F5 even when a response to that object is not required. The possible visual or motor nature of this object coding is discussed.

Animals↗

Premotor cortex and the recognition of motor actions.

In area F5 of the monkey premotor cortex there are neurons that discharge both when the monkey performs an action and when he observes a similar action made by another monkey or by the experimenter. We report here some of the properties of these 'mirror' neurons and we propose that their activity 'represents' the observed action. We posit, then, that this motor representation is at the basis of the understanding of motor events. Finally, on the basis of some recent data showing that, in man, the observation of motor actions activate the posterior part of inferior frontal gyrus, we suggest that the development of the lateral verbal communication system in man derives from a more ancient communication system based on recognition of hand and face gestures.

Animals↗

Action recognition in the premotor cortex.

We recorded electrical activity from 532 neurons in the rostral part of inferior area 6 (area F5) of two macaque monkeys. Previous data had shown that neurons of this area discharge during goal-directed hand and mouth movements. We describe here the properties of a newly discovered set of F5 neurons ("mirror neurons', n = 92) all of which became active both when the monkey performed a given action and when it observed a similar action performed by the experimenter. Mirror neurons, in order to be visually triggered, required an interaction between the agent of the action and the object of it. The sight of the agent alone or of the object alone (three-dimensional objects, food) were ineffective. Hand and the mouth were by far the most effective agents. The actions most represented among those activating mirror neurons were grasping, manipulating and placing. In most mirror neurons (92%) there was a clear relation between the visual action they responded to and the motor response they coded. In approximately 30% of mirror neurons the congruence was very strict and the effective observed and executed actions corresponded both in terms of general action (e.g. grasping) and in terms of the way in which that action was executed (e.g. precision grip). We conclude by proposing that mirror neurons form a system for matching observation and execution of motor actions. We discuss the possible role of this system in action recognition and, given the proposed homology between F5 and human Brocca's region, we posit that a matching system, similar to that of mirror neurons exists in humans and could be involved in recognition of actions as well as phonetic gestures.

Animals↗

Coding of peripersonal space in inferior premotor cortex (area F4).

1. We studied the functional properties of neurons in the caudal part of inferior area 6 (area F4) in awake monkeys. In agreement with previous reports, we found that the large majority (87%) of neurons responded to sensory stimuli. The responsive neurons fell into three categories: somatosensory neurons (30%); visual neurons (14%); and bimodal, visual and somatosensory neurons (56%). Both somatosensory and bimodal neurons typically responded to light touch of the skin. Their RFs were located on the face, neck, trunk, and arms. Approaching objects were the most effective visual stimuli. Visual RFs were mostly located in the space near the monkey (peripersonal space). Typically they extended in the space adjacent to the tactile RFs. 2. The coordinate system in which visual RFs were coded was studied in 110 neurons. In 94 neurons the RF location was independent of eye position, remaining in the same position in the peripersonal space regardless of eye deviation. The RF location with respect to the monkey was not modified by changing monkey position in the recording room. In 10 neurons the RF's location followed the eye movements, remaining in the same retinal position (retinocentric RFs). For the remaining six neurons the RF organization was not clear. We will refer to F4 neurons with RF independent of eye position as somatocentered neurons. 3. In most somatocentered neurons (43 of 60 neurons) the background level of activity and the response to visual stimuli were not modified by changes in eye position, whereas they were modulated in the remaining 17. It is important to note that eye deviations were constantly accompanied by a synergic increase of the activity of the ipsilateral neck muscles. It is not clear, therefore, whether the modulation of neuron discharge depended on eye position or was a consequence of changes in neck muscle activity. 4. The effect of stimulus velocity (20-80 cm/s) on neuron response intensity and RF extent in depth was studied in 34 somatocentered neurons. The results showed that in most neurons the increase of stimulus velocity produced an expansion in depth of the RF. 5. We conclude that space is coded differently in areas that control somatic and eye movements. We suggest that space coding in different cortical areas depends on the computational necessity of the effectors they control.

Animals↗

Motor facilitation during action observation: a magnetic stimulation study.

1. We stimulated the motor cortex of normal subjects (transcranial magnetic stimulation) while they 1) observed an experimenter grasping 3D-objects, 2) looked at the same 3D-objects, 3) observed an experimenter tracing geometrical figures in the air with his arm, and 4) detected the dimming of a light. Motor evoked potentials (MEPs) were recorded from hand muscles. 2. We found that MEPs significantly increased during the conditions in which subjects observed movements. The MEP pattern reflected the pattern of muscle activity recorded when the subjects executed the observed actions. 3. We conclude that in humans there is a system matching action observation and execution. This system resembles the one recently described in the monkey.

Arm↗