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Biomedical subjects

G Rizzolatti

Publications and source records attributed to G Rizzolatti.

At least 19 recordsLinked to original sources

Space coding by premotor cortex.

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.

Animals

Understanding motor events: a neurophysiological study.

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.

Animals

Motor control of voluntary arm movements. Kinematic and modelling study.

The motor control of pointing and reaching-to-grasp movements was investigated using two different approaches (kinematic and modelling) in order to establish whether the type of control varies according to modifications of arm kinematics. Kinematic analysis of arm movements was performed on subjects' hand trajectories directed to large and small stimuli located at two different distances. The subjects were required either to grasp and to point to each stimulus. The kinematics of the subsequent movement, during which subject's hand came back to the starting position, were also studied. For both movements, kinematic analysis was performed on hand linear trajectories as well as on joint angular trajectories of shoulder and elbow. The second approach consisted in the parametric identification of the black box (ARMAX) model of the controller driving the arm movement. Such controller is hypothesized to work for the correct execution of the motor act. The order of the controller ARMAX model was analyzed with respect to the different experimental conditions (distal task, stimulus size and distance). Results from kinematic analysis showed that target distance and size influenced kinematic parameters both of angular and linear displacements. Nevertheless, the structure of the motor program was found to remain constant with distance and distal task, while it varied with precision requirements due to stimulus size. The estimated model order of the controller confirmed the invariance of the control law with regard to movement amplitude, whereas it was sensitive to target size.

Adult

Architecture of superior and mesial area 6 and the adjacent cingulate cortex in the macaque monkey.

The agranular frontal cortex is formed by several distinct functional areas. There is no agreement, however, on its cytoarchitectonic organization. The aim of this study was to redefine the cytoarchitectonic organization of superior and mesial area 6 and the adjacent cingulate cortex in the macaque monkey. A particular goal was to find out whether the so-called supplementary motor area (SMA) is cytoarchitectonically different from the rest of area 6 and whether it can be considered as a single, independent cytoarchitectonic area. The results showed that, rostral to F1 (area 4), four architectonic areas can be recognized in the superior (dorsal) and mesial area 6. Two fo them are located on mesial cortical surface (F3 caudally and F6 rostrally) and two on superior cortical convexity (F2 caudally and F7 rostrally). The main cytoarchitectonic features of the five identified areas can be summarized as follows. F1: (1) giant pyramidal cells organized in multiple rows, (2) columnar pattern extending from the white matter to the superficial layers, (3) low cellular density in the lower part of layer III. F3: (1) high cellular density in the lower part of layer III, which fuses with a dense Va, (2) columnar pattern present only in the deepest layer, (3) occasional presence of giant pyramidal cells in layer Vb. F6: (1) prominent layer V, (2) absence of sublayer Vb, (3) homogeneous cell density in superficial layers. F2: (1) thin row of medium-size pyramids in the lowest part of layer III, (2) columnar pattern extending to the superficial layers, (3) dense layer Va, (4) few, scattered giant pyramids in layer Vb. F7: (1) prominent layer V, (2) bipartite layer VI. Areas F1, F2, and F3, as defined cytoarchitectonically, coincided with the homonymous histochemical areas. The present data showed also that area 24 is formed by four subareas: 24a, b, c and d. Areas 24a and b occupy the ventral part of area 24, whereas its dorsal part is formed by area 24c, located rostrally, and area 24d, located caudally. The following features distinguish area 24d from area 24c: (1) larger pyramidal cells in layer V, (2) presence of medium-size pyramidal cells in the lower part of layer III, (3) more prominent columnar pattern, (4) higher myelinization with the presence of an evident horizontal plexus. Mesial area 6 is usually considered as a single functional entity (SMA). Our findings show that this cortical region is formed by two distinct cytoarchitectonic areas.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Multiple representations of body movements in mesial area 6 and the adjacent cingulate cortex: an intracortical microstimulation study in the macaque monkey.

The mesial agranular frontal cortex that lies rostral to area 4 (F1) is formed by two distinct cytoarchitectonic areas: F3, located caudally, and F6, located rostrally. In the present experiments we investigated the organization of F3 and F6 by observing the motor responses evoked by their intracortical electrical microstimulation. Our main purpose was to find out whether the cytoarchitectonic subdivision of the mesial agranular frontal cortex into two areas has a physiological counterpart. The result showed that F3 (the caudal area) contains a complete motor representation with hindlimb movements located caudally, forelimb movements located centrally, and orofacial movements located rostrally. The great majority of limb movements involved proximal joints. With respect to F1, F3 showed the following functional characteristics: (1) lack of segregation between proximal and distal movements, (2) larger percentage of complex movements, and (3) higher excitability threshold. Movements were more difficult to elicit from F6 (the rostral area) than from F3. However, by using a longer stimulus train duration (100 ms) 39.3% of tested sites produced body movements. This percentage increased (50.5%) when the electrical stimulation was applied during monkey natural movements instead of when the monkey was still in its chair. Most of the evoked movements concerned the forelimb. More rarely, neck and upper face movements were observed. Unlike F1 and F3 where most movements were fast, slow movements were frequently observed with stimulation of F6. Many of them mimicked natural movements of the animal. Eye movements were evoked from F7 (superior area 6) but not from F6. An additional motor representation was found in the dorsocaudal part of area 24 (24d). This area is topographically organized with a forelimb representation located caudally and ventrally and a hindlimb representation located rostrally and dorsally. The excitability threshold of area 24d is higher than that of F1 and F3. Evoked movements were occasionally observed also after stimulation of area 24c. In conclusion, on the mesial cortical wall rostral to F1, there are at least three independent motor representations. On the basis of somatotopic organization and excitability properties, we propose that the term supplementary motor area (SMA-proper) should be reserved to F3.

Animals

Differential hemispheric asymmetries in depression and anxiety: a reaction-time study.

Several lines of evidence suggest impaired right hemisphere function in depression. In order to further investigate this phenomenon, simple reaction times (RTs) to lateralized visual stimuli were studied in patients with mild unipolar depression, in patients with chronic anxiety, and in medical patients free of psychiatric symptoms. The results showed a marked slowing of RTs to left visual field (right hemisphere) stimuli in depressed patients. Anxious patients showed a trend toward an opposite asymmetry, with slower responses to right field (left hemisphere) stimuli. In the nonpsychiatric group no visual field differences were present. The results are discussed in light of two alternative interpretations: depression may engage the right hemisphere's mechanisms, interfering with its functioning at a premotor level, or it may influence the regulation of performance by arousal and vigilance mechanisms lateralized to the right hemisphere, possibly operating at an earlier sensory stage.

Adult

Influence of different types of grasping on the transport component of prehension movements.

The main aim of the present study was to clarify whether different types of grasping may affect the transport component of prehension movements. To this purpose two experiments were carried out. In the first experiment the kinematics of arm movements (transport and manipulation components) were studied in eight normal subjects instructed to reach and grasp different objects located either 20 or 30 cm from their hand. The objects employed required two different types of grip: prehension with the whole hand and prehension with the index finger and the thumb (precision grip). In the second experiment subjects were instructed to point to the same objects employed in the first experiment. This experiment served as a control for the precision requirements related to the object size. The results showed that, once the precision requirements were taken into account, the transport component remained unmodified with the different types of grip. The time course of the manipulation component and its temporal relations with the transport component changed with the type of grasping. The maximal hand aperture was reached earlier in the precision grip than in the whole hand prehension and the temporal coupling with the transport component was weaker in the former condition than in the latter. The data are interpreted as further evidence in favour of independence between the transport and the manipulation "channels".

Adolescent

[Orientation of attention in the visual space].

The display was composed of four boxes, horizontally aligned above the fixation point. In Experiment I, each box was cued by a digit shown at fixation. In Experiment II there were only two numeric cues, signalling the inner or the outer boxes, depending on the experimental condition. The subject was instructed to orient attention to the cued box, and to respond to the imperative stimulus as fast as possible, wherever it appeared. By using four time interval (SOAs), we tried to determine the route covered by attention movements. In Experiment I, with the shortest SOA (100 msec), it was shown that attention does not reach the cued box through a direct path. Rather it moves first on the inner boxes, thereafter focusing on the cued location. The same results were obtained in Experiment II, where the cue directed attention to the inner boxes. When the external boxes were cued, however, this trend was not observed.

Adult

[Orientation strategy implicit in spatial attention].

In this experiment, two rows of four boxes, one above and the other below the fixation point, were displayed on a computer monitor. A cognitive numeric cue indicated one of the four external boxes, where the attention was to be directed. Stimuli, however, could occur in any of the eight boxes. The subject was instructed to respond to the imperative stimulus as fast as possible, regardless of its location. Four time intervals between cue and stimulus presentation (SOAs) were employed in order to disclose the route along which attention is moved. With the shortest SOA (100 msec) an indication that attention movements follow the cartesian/orthogonal axes was shown.

Adult

[Kinematic study of reaching-grasping movements in the monkey].

Kinematics of reaching-grasping movement towards stimuli of three different sizes located at two different distances were studied in one monkey (Macaca nemestrina). Transport and manipulation components were analyzed using the ELITE system. Transport time, peak velocity and deceleration phase of velocity were influenced by stimulus size, whilst acceleration phase remained unmodified. Peak velocity clearly increased with distance, while transport time remained constant (isochrony ). The main parameters of manipulation component were all influenced by stimulus size but they did not vary with distance. A comparison with kinematic data obtained from human subjects was made.

Animals

Cortico-cortical connections of two electrophysiologically identified arm representations in the mesial agranular frontal cortex.

Neuronal tracers (diamidino yellow or wheat germ agglutinin conjugated with horseradish peroxidase) were injected in the arm representations of area 6a alpha (mesial surface, area F3), in the arm representation of area 6a beta (mesial surface) as well as in the eye field of area 6a beta (dorso-medial surface). The results showed that the arm representation of area F3 receives topographically organized afferents from motor and premotor areas (areas F1, F2, F4 and F5). A further connection was found with that part of cingulate cortex that sends projections to the spinal cord. In contrast, the arm representation of area 6a beta receives afferents chiefly from area F5, the prefrontal cortex and that part of cingulate sulcus which has few, if any, connections with the spinal cord. No connections were found with the precentral motor cortex (area F1). The area 6a beta eye field receives afferents mostly from the frontal eye field. Further connections are with the prefrontal cortex and cingulate gyrus. It is suggested that the so called "low level" motor functions of supplementary motor area are due to the activity of area F3, whereas the so called "high level" motor functions depend upon an independent area located in area 6a beta.

Animals

Neurons related to reaching-grasping arm movements in the rostral part of area 6 (area 6a beta).

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.

Animals

Neglect as a neural representation deficit.

In recent years two competing sets of theories have been advanced in order to explain neglect: the attentional theories and the representational theories. In this article we examine to what extent these explanations of neglect are consistent with the neurophysiological data. After a brief review of some crucial works on animals and man related to neglect, we conclude that this syndrome is essentially a representational deficit consequent to lesions of neural centers responsible for the organization of motor acts, in which space is coded in non-retinal coordinates. The attentional disturbances are interpreted as a secondary factor which may aggravate the syndrome.

Animals

Thalamic input to inferior area 6 and area 4 in the macaque monkey.

Recent cytoarchitectonic, histochemical, and hodological studies in primates have shown that area 6 is formed by three main sectors: the supplementary motor area, superior area 6, which lies medial to the spur of the arcuate sulcus, and inferior area 6, which is located lateral to it. Inferior area 6 has been further subdivided into two histochemical areas: area F5, located along the inferior limb of the arcuate sulcus, and area F4, located between area F5 and area 4 (area F1). The present study traced the thalamocortical projections of inferior area 6 and the adjacent part of area 4 by injecting small amounts of WGA-HRP in specific sectors of the agranular frontal cortex. Our data showed that each histochemical area receives a large projection from one nucleus of the ventrolateral thalamus (motor thalamus) and additional projections from other nuclei of this thalamic sector. Area F5 receives a large projection from area X of Olszewski ('52) and additional projections from the caudal part of the nucleus ventralis posterior lateralis, pars oralis (VPLo), and the nucleus ventralis lateralis, pars caudalis (VLc) (VPLo-VLc complex). Area F4 receives a large projection from the nucleus ventralis lateralis, pars oralis (VLo), and additional projections from area X and the VPLo-VLc complex. The rostral part of area F1 is innervated chiefly by VLo, plus smaller contributions from rostral VPLo and the VPLo-VLc complex. The caudal part of F1 receives its greatest input from VPLo, with a small contribution from VLo. In addition, each histochemical area receives projections originating from the intralaminar thalamic nuclei, the posterior thalamus, and--for area F4 and area F5--also from the nucleus medialis dorsalis (MD). Analysis of the physiological properties of the various histochemical areas in relation to their main thalamic input showed that those cortical fields in which distal movements are predominant (area F5, caudal part of area F1) are innervated chiefly by area X and VPLo, whereas those cortical fields in which proximal movements are predominant receive their main input from VLo. Because VPLo and area X are targets of cerebellothalamic pathways, whereas VLo receives a pallidal input, we propose that the cortical fields in which distal movements are most heavily represented are mainly under the influence of the cerebellum, whereas the cortical fields in which proximal movements are most heavily represented are mainly under the influence of the basal ganglia.

Animals

Somatotopic representation in inferior area 6 of the macaque monkey.

On the basis of its cytoarchitectonic and enzymatic properties area 6 of the macaque monkey can be subdivided into two large sectors: a superior sector lying medial to the spur of the arcuate sulcus (superior area 6 or F2) and an inferior sector lying lateral to it (inferior area 6). Inferior area 6 is constituted by two enzymatic areas: F4 and F5. In this study we investigated the somatotopic organization of inferior area 6 and the adjacent area 4 combining single-neuron recording and intracortical electrical microstimulation. We found that two separate movement representations exist in this region. The caudal one corresponds to area F1 (primary motor cortex), the rostral one to inferior area 6. The two representations are mirror images one of the other with the axioproximal movements being adjacently located. In the rostral map the proximal movements are mostly located in F4, the distal movements in F5. Neuronal properties indicate that the rostral map has characteristics that are more complex than the caudal map. We propose that the rostral map is involved in transforming visual information in motor commands. F4 should be involved in the control of arm movements based on the location of the objects in respect to the body, whereas F5 should play a role in the control of grasping movements on the basis of the size of the stimuli.

Animals

[Kinematic study of reaching-grasping movements].

The experiment was conducted to investigate, by using kinematic parameters, the influence of the type of prehension on the transportation component in reaching-grasping movements. The main question was whether the transportation component is influenced by the type of prehension besides the distance of the object. The experiment was carried out on eight subjects who performed reaching-grasping movements toward objects located at different distances. Two types of prehension were examined: whole hand prehension and precision grip. The following kinematic parameters of the transportation component (wrist movement) were studied: movement times, profiles of velocity and accelerations. Our results have shown that the transportation component is affected by the two factors. However the kinematic parameters were influenced differently by the distance and the type of prehension. Our conclusion is that, although distance and type of prehension affect the transportation component, they are computed separately in programming this component.

Adolescent

Functional organization of inferior area 6 in the macaque monkey. I. Somatotopy and the control of proximal movements.

Two series of experiments are reported in this paper. The first concerns the movement representation in the macaque inferior area 6, the second the functional properties of neurons located in the caudal part of this area (histochemical area F4). By combining single neuron recording and intracortical microstimulation, we found that inferior area 6 is somatotopically organized. The axio-proximal movements are represented caudally, the distal movements are represented near the arcuate sulcus. The mouth field is located laterally, the hand field medially. There is no leg field. A comparison between neuron properties and histochemical characteristics of inferior area 6 showed that the proximal movements representation includes most of area F4, whereas the distal movements representation corresponds to area F5 and to the rostral part of F4. Neurons located in that part of F4 where proximal movements are represented respond very well to tactile stimuli. They have large receptive fields mostly located on the face and on the upper part of the body. A large number of these neurons respond to visual stimuli. Objects approaching the animal are particularly effective. The tactile and the visual receptive fields are in register. The most represented movements are reaching movements, movements bringing the hand to the mouth or to the body and facial movements. There is a congruence between location of visual fields and preferred arm movements. It is argued that the receptive field arrangement and the response properties are more complex in area F4 than in the primary motor cortex and that area F4 neurons are involved in the control of arm movements towards different space sectors.

Animals