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MOTOR CONTROL SERIESThe Premotor Cortexand Mirror NeuronsIn 1991, Nature rejected the first report on mirror neurons for its 'lack of general interest'. Undeterred,the research team managed to publish a report the following year, and mirror neurons have been in thenews ever since. Indeed, claims that mirror neurons underpin such functions as language acquisition,theory of mind and empathy have been made. Here, Fogassi (one of the authors of the original report)and Rodà present an account of mirror neurons and motor control.Martyn Bracewell, Series editorLeonardo Fogassiis full professor of Physiology atthe Department of Psychology ofthe University of Parma. He has afirst degree (cum laude) in Biologyand a PhD in Neuroscience. Hismain research interests concernthe neural cortical mechanisms ofsensorimotor transformation andthe cognitive properties of themotor system in monkeys andhumans, with particular focus onthe mirror neuron system.Francesca RodàPsychologist, PhD in Neuroscience,is a post-doc at The Department ofNeuroscience, Section of HumanPhysiology, of the University ofParma. Her research interestsinclude investigations of monkeymirror neurons and their role inaction and intention understandingand of the neural bases of communication/languageevolution.Several neurophysiological studies in monkeysdemonstrated that neurons of the agranularfrontal cortex code goal-related motor acts,such as reaching an object, grasping it, etc., ratherthan simple movements. In particular, single neuronsof ventral premotor area F5 (Figure 1) code themotor goal at an abstract level, discharging when amonkey grasps an object independent of whetherthis act is performed with the hand, the mouth oreven with a tool. 1 This “internal motor knowledge” isthen exploited, through reciprocal anatomicalconnections between parietal and premotor cortex,by the incoming sensory information, constituting asystem matching the sensory input onto specificmotor representations. This system enables individualsto attribute a “motor meaning” to the sensoryinput. One of the best examples of this matchingprocess is provided by mirror neurons.Mirror neurons in the monkeyMirror neurons, originally described in monkey areaF5, are visuomotor neurons discharging both when amonkey performs a hand or mouth goal-directedmotor act (e.g. grasping, biting, or manipulating anobject) and when it observes the same or a similaract performed by another individual (Figure 2). Asub-class of mirror neurons respond not only duringexecution and observation of a motor act, but alsoto the sound of noisy motor acts such as peanutbreaking. 1 Although mirror neurons are generallynot influenced by many details of the observedmotor acts, recently it has been demonstrated that aconsistent number of them can be modulated bythe visual perspective (egocentric or third personview) from which a motor act is observed 2 or by thedistance at which the observed act is performed. 3Thus these neurons, beyond encoding the goal ofthe observed motor acts, can also contribute torecognize some details of it, probably through feedbackconnections between ventral premotor cortexand posterior, high order visual areas.The idea that mirror neurons have a crucial role inthe understanding of motor acts has been supportedby further neurophysiological investigations. In one ofthese 4 it has been demonstrated that mirror neuronsdischarge also when the hand-target interaction ishidden behind a screen, thus showing that the motorrepresentation of the observed motor act is retrievedeven in absence of its full visual description.The presence of mirror neurons has been demonstratedalso in the inferior parietal cortex, in a cytoarchitectonicarea (PFG) strictly linked with the F5“mirror” sector. Thus, these two areas, together withSTS (containing visual neurons responding to theCorrespondence to:Leonardo Fogassi,Dipartimento di Neuroscienze,Università di Parma,via Volturno 39, 43125 - Parma, Italy.Tel: +39 521 903847Fax: +39 521 903900Email: leonardo.fogassi@unipr.itFigure 1: Lateral View of the monkey brain showing the parcellation of the agranular frontal and posterior parietal cortices. Motorareas are indicated with the letter F followed by a number. The areas forming the posterior parietal cortex are indicated with theletter P, followed by another letter, except the most posterior part of the inferior parietal cortex (Opt). Abbreviations: AI, inferiorarcuate sulcus; AS, superior arcuate sulcus; C, central sulcus; IP, intraparietal sulcus; L, lateral fissure;P, principal sulcus; STS, superior temporal sulcus.<strong>ACNR</strong> > VOLUME 11 NUMBER 4 > SEPTEMBER/OCTOBER 2011 > 23

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