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Many neurons in the monkey ventral premotor area F5 discharge selectively when the monkey grasps an object with a specific grip. Of these, the motor neurons are active only during grasping execution, whereas the visuomotor neurons also respond to object presentation. Here we assessed whether the activity of 90 task-related F5 neurons recorded from two macaque monkeys during the performance of a visually-guided grasping task can be used as input to pattern recognition algorithms aiming to decode different grips. The features exploited for the decoding were the mean firing rate and the mean interspike interval calculated over different time spans of the movement period (all neurons) or of the object presentation period (visuomotor neurons). A support vector machine (SVM) algorithm was applied to the neural activity recorded while the monkey grasped two sets of objects. The original set contained three objects that were grasped with different hand shapes, plus three others that were grasped with the same grip, whereas the six objects of the special set were grasped with six distinctive hand configurations. The algorithm predicted with accuracy greater than 95% all the distinct grips used to grasp the objects. The classification rate obtained using the first 25% of the movement period was 90%, whereas it was nearly perfect using the entire period. At least 16 neurons were needed for accurate performance, with a progressive increase in accuracy as more neurons were included. Classification errors revealed by confusion matrices were found to reflect similarities of hand grips used to grasp the objects. The use of visuomotor neurons' responses to object presentation yielded grip classification accuracy similar to that obtained from actual grasping execution. We suggest that F5 grasping-related activity might be used by neural prostheses to tailor hand shape to the specific object to be grasped even before movement onset., (Copyright © 2011 IBRO. Published by Elsevier Ltd. All rights reserved.)

The capacity to use tools is a fundamental evolutionary achievement. Its essence stands in the capacity to transfer a proximal goal (grasp a tool) to a distal goal (e.g., grasp food). Where and how does this goal transfer occur? Here, we show that, in monkeys trained to use tools, cortical motor neurons, active during hand grasping, also become active during grasping with pliers, as if the pliers were now the hand fingers. This motor embodiment occurs both for normal pliers and for "reverse pliers," an implement that requires finger opening, instead of their closing, to grasp an object. We conclude that the capacity to use tools is based on an inherently goal-centered functional organization of primate cortical motor areas.

A direct form of 'experiential understanding' of others is achieved by modelling their behaviours as intentional experiences on the basis of the equivalence between what the others do and feel and what we do and feel. This modelling mechanism is embodied simulation. By means of embodied simulation we do not just 'see' an action, an emotion, or a sensation. Side by side with the sensory description of the observed social stimuli, internal representations of the body states associated with actions, emotions, and sensations are evoked in the observer, as if he/she would be doing a similar action or experiencing a similar emotion or sensation. Mirror neurons are likely the neural correlate of this mechanism. The mirror neuron matching systems map the different intentional relations in a compressed fashion, which is neutral about the specific quality or identity of the agentive/subjective parameter. By means of a shared neural state realized in two different bodies that nevertheless obey to the same functional rules, the 'objectual other' becomes 'another self'.

Understanding the intentions of others while watching their actions is a fundamental building block of social behavior. The neural and functional mechanisms underlying this ability are still poorly understood. To investigate these mechanisms we used functional magnetic resonance imaging. Twenty-three subjects watched three kinds of stimuli: grasping hand actions without a context, context only (scenes containing objects), and grasping hand actions performed in two different contexts. In the latter condition the context suggested the intention associated with the grasping action (either drinking or cleaning). Actions embedded in contexts, compared with the other two conditions, yielded a significant signal increase in the posterior part of the inferior frontal gyrus and the adjacent sector of the ventral premotor cortex where hand actions are represented. Thus, premotor mirror neuron areas-areas active during the execution and the observation of an action-previously thought to be involved only in action recognition are actually also involved in understanding the intentions of others. To ascribe an intention is to infer a forthcoming new goal, and this is an operation that the motor system does automatically.

We investigated the properties of neurons located in the distal forelimb field of dorsal premotor area F2 of macaque monkey using a behavioral paradigm for studying the neuronal discharge during observation (object fixation condition) and grasping of different 3-dimensional objects with and without visual guidance of the movement (movement in light and movement in dark conditions, respectively). The main result is that almost all studied neurons were selective for both the type of prehension and the wrist orientation required for grasping an object. Three categories of neurons were found: purely motor, visually modulated, and visuomotor neurons. The discharge of purely motor neurons was not affected by either object presentation or by the visual feedback of the hand approaching to and interacting with the object. Visually modulated neurons presented a different discharge in the 2 movement conditions, this determining a decrease in selectivity for the grip and wrist orientation in the movement in dark condition. Visuomotor neurons typically discharged during the object fixation task even in the absence of any grasping movement. Nine of them also displayed a different discharge rate between the 2 movement conditions. Congruence was observed between the neuron response during the most effective type of prehension and the neuron response during observation of the object requiring that particular prehension. These results indicate an important role of F2 in the control of goal-related hand movements.

Starting from a neurobiological standpoint, I will propose that our capacity to understand others as intentional agents, far from being exclusively dependent upon mentalistic/linguistic abilities, be deeply grounded in the relational nature of our interactions with the world. According to this hypothesis, an implicit, prereflexive form of understanding of other individuals is based on the strong sense of identity binding us to them. We share with our conspecifics a multiplicity of states that include actions, sensations and emotions. A new conceptual tool able to capture the richness of the experiences we share with others will be introduced: the shared manifold of intersubjectivity. I will posit that it is through this shared manifold that it is possible for us to recognize other human beings as similar to us. It is just because of this shared manifold that intersubjective communication and ascription of intentionality become possible. It will be argued that the same neural structures that are involved in processing and controlling executed actions, felt sensations and emotions are also active when the same actions, sensations and emotions are to be detected in others. It therefore appears that a whole range of different "mirror matching mechanisms" may be present in our brain. This matching mechanism, constituted by mirror neurons originally discovered and described in the domain of action, could well be a basic organizational feature of our brain, enabling our rich and diversified intersubjective experiences. This perspective is in a position to offer a global approach to the understanding of the vulnerability to major psychoses such as schizophrenia., (Copyright 2003 S. Karger AG, Basel)

In the ventral premotor cortex (area F5) of the monkey there are neurons that discharge both when the monkey performs specific motor actions and when it observes another individual performing a similar action (mirror neurons). Previous studies on mirror neurons concerned hand actions. Here, we describe the mirror responses of F5 neurons that motorically code mouth actions. The results showed that about one-third of mouth motor neurons also discharge when the monkey observes another individual performing mouth actions. The majority of these 'mouth mirror neurons' become active during the execution and observation of mouth actions related to ingestive functions such as grasping, sucking or breaking food. Another population of mouth mirror neurons also discharges during the execution of ingestive actions, but the most effective visual stimuli in triggering them are communicative mouth gestures (e.g. lip smacking). Some also fire when the monkey makes communicative gestures. These findings extend the notion of mirror system from hand to mouth action and suggest that area F5, the area considered to be the homologue of human Broca's area, is also involved in communicative functions.

Many object-related actions can be recognized by their sound. We found neurons in monkey premotor cortex that discharge when the animal performs a specific action and when it hears the related sound. Most of the neurons also discharge when the monkey observes the same action. These audiovisual mirror neurons code actions independently of whether these actions are performed, heard, or seen. This discovery in the monkey homolog of Broca's area might shed light on the origin of language: audiovisual mirror neurons code abstract contents-the meaning of actions-and have the auditory access typical of human language to these contents.

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.

In this study, we mainly investigated the visual selectivity of hand-manipulation-related neurons in the anterior intraparietal area (area AIP) while the animal was grasping or fixating on three-dimensional (3D) objects of different geometric shapes, sizes, and orientations. We studied the activity of 132 task-related neurons during the hand-manipulation tasks in the light and in the dark, as well as during object fixation. Seventy-seven percent (101/132) of the hand-manipulation-related neurons were visually responsive, showing either lesser activity during manipulation in the dark than during that in the light (visual-motor neurons) or no activation in the dark (visual-dominant neurons). Of these visually responsive neurons, more than half (n = 66) responded during the object-fixation task (object-type). Among these, 55 were tested for their shape selectivity during the object-fixation task, and many (n = 25) were highly selective, preferring one particular shape of the six different shapes presented (ring, cube, cylinder, cone, sphere, and square plate). For 28 moderately selective object-type neurons, we performed multidimensional scaling (MDS) to examine how the neurons encode the similarity of objects. The results suggest that some moderately selective neurons responded preferentially to common geometric features shared by similar objects (flat, round, elongated, etc.). Moderately selective nonobject-type visually responsive neurons, which did not respond during object fixation, were found by MDS to be more closely related to the handgrip than to the object shape. We found a similar selectivity for handgrip in motor-dominant neurons that did not show any visual response. With regard to the size of the objects, 16 of 26 object-type neurons tested were selective for both size and shape, whereas 9 object-type neurons were selective for shape but not for size. Seven of 12 nonobject-type and all (8/8) of the motor-dominant neurons examined were selective for size, and almost all of them were also selective for objects. Many hand-manipulation-related neurons that preferred the plate and/or ring were selective for the orientation of the objects (17/20). These results suggest that the visual responses of object-type neurons represent the shape, size, and/or orientation of 3D objects, whereas those of the nonobject-type neurons probably represent the shape of the handgrip, grip size, or hand-orientation. The activity of motor-dominant neurons was also, in part, likely to represent these parameters of hand movement. This suggests that the dorsal visual pathway is concerned with the aspect of form, orientation, and/or size perception that is relevant for the visual control of movements.

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.

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.

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.

Neurons of the rostral part of inferior premotor cortex of the monkey discharge during goal-directed hand movements such as grasping, holding, and tearing. We report here that many of these neurons become active also when the monkey observes specific, meaningful hand movements performed by the experimenters. The effective experimenters' movements include among others placing or retrieving a piece of food from a table, grasping food from another experimenter's hand, and manipulating objects. There is always a clear link between the effective observed movement and that executed by the monkey and, often, only movements of the experimenter identical to those controlled by a given neuron are able to activate it. These findings indicate that premotor neurons can retrieve movements not only on the basis of stimulus characteristics, as previously described, but also on the basis of the meaning of the observed actions.

Many neurons in inferior area 6, a cortical premotor area, respond to visual stimuli presented in the space around the animal. We were interested to learn whether the receptive fields of these neurons are coded in retinotopic or in body-centered coordinates. To this purpose we recorded single neurons from inferior area 6 (F4 sector) in a monkey trained to fixate a light and detect its dimming. During fixation visual stimuli were moved towards the monkey both within and outside the neuron's receptive field. The fixation point was then moved and the neuron retested with the monkey's gaze deviated to the new location. The results showed that most inferior area 6 visual neurons code the stimulus position in spatial and not in retinal coordinates. It is proposed that these visual neurons are involved in generating the stable body-centered frame of reference necessary for programming visually guided movements.

Single neurons were recorded from the rostral part of the agranular frontal cortex (area 6a beta) in awake, partially restrained macaque monkeys. In the medialmost and mesial sectors of this area, rostral to the supplementary motor area, neurons were found which were activated during arm reaching-grasping movements. These neurons ("reaching-grasping neurons") did not appear to be influenced by how the objects were grasped nor, with some exceptions, by where they were located. Their activity changed largely prior to the arm movement and continued until the end of it. The premovement modulation (excitatory or inhibitory) could start with stimulus presentation, with the saccade triggered by the stimulus or after stimulus fixation. The distance of the stimulus from the monkey was an important variable for activating many neurons. About half of the recorded neurons showed a modulation of the same sign during movement and premovement period. The other half showed an increase/decrease in activity which was of the opposite sign during movement and premovement period or part of it. In this last case the discharge changes were of the same sign when the stimulus was close to the monkey and when the monkey moved its arm to reach the objects, whereas they were of opposite sign when the stimulus was outside the animal's reach. Microstimulation of area 6a beta and the reconstruction of the locations of eye movement and arm movement related cells showed that the arm field was located more medially (and mesially) than the eye field described by Schlag and Schlag-Rey (1987). It is suggested that, unlike inferior area 6, which is mostly involved in selection of effectors on the basis of the physical properties of the objects and their spatial location (Rizzolatti and Gentilucci 1988), area 6a beta plays a role in the preparation of reaching-grasping arm movements and in their release when the appropriate conditions are set.

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. [ABSTRACT FROM AUTHOR]

Our seemingly effortless capacity of conceiving of the acting bodies inhabiting our social world as goal-oriented individuals like us depends on the constitution of a shared “we-centric” space. I have proposed that this shared manifold space can be characterized at the functional level as embodied simulation, a basic functional mechanism by means of which our brain/body system models its interactions with the world. The mirroring mechanism for action and other mirroring mechanisms in our brain represent sub-personal instantiations of embodied simulation. Embodied simulation provides a new empirically based notion of intersubjectivity, viewed first and foremost as intercorporeity. Embodied simulation challenges the notion that Folk-psychology is the sole account of interpersonal understanding. Before and below mind reading is intercorporeity as the main source of knowledge we directly gather about others. By means of embodied simulation we can map others’ actions onto our own motor representations, as well as others’ emotions and sensations onto our own viscero-motor and somatosensory representations. “Representation”, as used here, refers to a particular type of content, generated by the relations that our situated and inter-acting brain-body system instantiates with the world. Such content is pre-linguistic and pre-theoretical, but nevertheless has attributes normally and uniquely attributed to conceptual content.Social cognition is not only explicitly reasoning about the contents of someone else''s mind. Embodied simulation, gives us a direct insight of other minds thus enabling our capacity to empathize with others. This proposal opens new perspectives on our understanding of autism and other psychopathological states such as schizophrenia. [Copyright &y& Elsevier]

The article cites a study which use functional magnetic resonance imaging to show that the mirror neuron system tracks not only the actions, but also the intentions, of others. The study was conducted by researcher Marco Iacoboni and colleagues. Mirror neurons are a set of neurons in the premotor area of the brain that are activated not only when performing an action oneself, but also while observing someone else perform that action. It is believed mirror neurons increase an individual's ability to understand the behaviors of others, an important skill in social species such as humans.