Luciano Fadiga

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 observers 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.

Localization of grasp representations in humans by PET: 1. Observation versus execution

Positron emission tomography (PET) was used to localize brain regions that are active during the observation of grasping movements. Normal, right-handed subjects were tested under three conditions. In the first, they observed grasping movements of common objects performed by the experimenter. In the second, they reached and grasped the same objects. These two conditions were compared with a third condition consisting of object observation. On the basis of monkey data, it was hypothesized that during grasping observation, activations should be present in the region of the superior temporal sulcus (STS) and in inferior area 6. The findings in humans demonstrated that grasp observation significantly activates the cortex of the middle temporal gyrus including that of the adjacent superior temporal sulcus (Brodmanns area 21) and the caudal part of the left inferior frontal gyrus (Brodmanns area 45). The possible functional homologies between these areas and the monkey STS region and frontal area F5 are discussed.

Localization of grasp representations in humans by positron emission tomography

Positron emission tomography imaging of cerebral blood flow was used to localize brain areas involved in the representation of hand grasping movements. Seven normal subjects were scanned under three conditions. In the first, they observed precision grasping of common objects performed by the examiner. In the second, they imagined themselves grasping the objects without actually moving the hand. These two tasks were compared with a control task of object viewing. Grasp observation activated the left rostral superior temporal sulcus, left inferior frontal cortex (area 45), left rostral inferior parietal cortex (area 40), the rostral part of left supplementary motor area (SMA-proper), and the right dorsal premotor cortex. Imagined grasping activated the left inferior frontal (area 44) and middle frontal cortex, left caudal inferior parietal cortex (area 40), a more extensive response in left rostral SMA-proper, and left dorsal premotor cortex. The two conditions activated different areas of the right posterior cerebellar cortex. We propose that the areas active during grasping observation may form a circuit for recognition of hand-object interactions, whereas the areas active during imagined grasping may be a putative human homologue of a circuit for hand grasping movements recently defined in nonhuman primates. The location of responses in SMA-proper confirms the rostrocaudal segregation of this area for imagined and real movement. A similar segregation is also present in the cerebellum, with imagined and observed grasping movements activating different parts of the posterior lobe and real movements activating the anterior lobe.

Premotor Cortex Activation during Observation and Naming of Familiar Tools

Positron emission tomography was used to investigate whether observation of real objects (tools of common use) activates premotor areas in the absence of any overt motor demand. Silent naming of the presented tools and silent naming of their use were also studied. Right-handed normal subjects were employed. Tool observation strongly activated the left dorsal premotor cortex. In contrast, silent tool naming activated Brocas area without additional activity in the dorsal premotor cortex. Silent tool-use naming, in addition to activating Brocas area, increased the activity in the left dorsal premotor cortex and recruited the left ventral premotor cortex and the left supplementary motor area. These data indicate that, even in the absence of any subsequent movement, the left premotor cortex processes objects that, like tools, have a motor valence. This dorsal premotor activation, which further augments when the subject names the tool use, should reflect the neural activity related to motor schemata for object use. The presence of an activation of both dorsal premotor cortex and ventral premotor cortex during tool-use naming suggests a role for these two areas in understanding object semantics.

Active perception: sensorimotor circuits as a cortical basis for language

Action and perception are functionally linked in the brain, but a hotly debated question is whether perception and comprehension of stimuli depend on motor circuits. Brain language mechanisms are ideal for addressing this question. Neuroimaging investigations have found specific motor activations when subjects understand speech sounds, word meanings and sentence structures. Moreover, studies involving transcranial magnetic stimulation and patients with lesions affecting inferior frontal regions of the brain have shown contributions of motor circuits to the comprehension of phonemes, semantic categories and grammar. These data show that language comprehension benefits from frontocentral action systems, indicating that action and perception circuits are interdependent.

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.

Hand action preparation influences the responses to hand pictures

The relations between stimuli triggering a hand grasping movement and the subsequent action were studied in normal human participants. Participants were instructed to prepare to grasp a bar, oriented either clockwise or counterclockwise, and to grasp it as fast as possible on presentation of a visual stimulus with their right hand. The visual stimuli were pictures of the right hand as seen in a mirror. In Experiment 1, they represented the mirror image of the hand final posture as achieved in grasping the bar oriented either clockwise or counterclockwise. In Experiment 2, in addition to the pictures of Experiment 1, another two pictures, obtained rotating the hands represented in the previous ones of 90 degrees, were also used. Both experiments showed that the reaction times were faster when there was a similarity between hand position as depicted in the triggering visual stimulus and the grasping hand final position, the fastest responses being those where this similarity was the closest. In addition, Experiment 2 showed that reaction times to not rotated stimuli were faster than reaction times to the rotated stimuli, thus excluding a simple stimulus-response compatibility explanation of the findings. The data are interpreted as behavioral evidence that there is a close link between specific visual stimuli and specific motor actions. A neurophysiological model for this visuo-motor link is presented.

The iCub humanoid robot: An open-systems platform for research in cognitive development

We describe a humanoid robot platform--the iCub--which was designed to support collaborative research in cognitive development through autonomous exploration and social interaction. The motivation for this effort is the conviction that significantly greater impact can be leveraged by adopting an open systems policy for software and hardware development. This creates the need for a robust humanoid robot that offers rich perceptuo-motor capabilities with many degrees of freedom, a cognitive capacity for learning and development, a software architecture that encourages reuse & easy integration, and a support infrastructure that fosters collaboration and sharing of resources. The iCub satisfies all of these needs in the guise of an open-system platform which is freely available and which has attracted a growing community of users and developers. To date, twenty iCubs each comprising approximately 5000 mechanical and electrical parts have been delivered to several research labs in Europe and to one in the USA.

The Imitative Mind: From mirror neurons to imitation: Facts and speculations

Introduction This chapter is composed of two parts. In the first we review the functional properties of an intriguing class of premotor neurons that we discovered in the monkey premotor cortex: the “mirror neurons.” These neurons discharge both when the monkey performs an action and when it observes another individual making a similar action. The second part is basically speculative. It is based on the hypothesis that there is a very general, evolutionary ancient mechanism, that we will name “resonance” mechanism, through which pictorial descriptions of motor behaviors are matched directly on the observers motor “representations” of the same behaviors. We will posit that resonance mechanism is a fundamental mechanism at the basis of inter-individual relations including some behaviors commonly described under the heading of “imitation.” Functional properties of area F5 Motor properties Area F5 forms the rostral part of inferior area 6 (Matelli et al ., 1985). Microstimulation and single-neuron studies showed that F5 contains a hand and a mouth movement representation (Gentilucci et al ., 1988; Hepp-Reymond, Husler, Maier, & Qi, 1994; Okano & Tanji, 1987; Rizzolatti et al ., 1981; Rizzolatti et al ., 1988). Particularly interesting results were obtained when F5 neurons were studied in a semi-naturalistic context (Rizzolatti et al ., 1988). Awake monkeys were seated on a primate chair and presented with various objects (geometrical solids, pieces of food of different size and shape). The stimuli were introduced in various spatial locations around the monkey, inside and outside its peripersonal space.

Space coding by premotor cortex

SummaryMany 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 neuronss receptive field. The fixation point was then moved and the neuron retested with the monkeys 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 necesary for programming visually guided movements.