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Biomedical subjects

G Rizzolatti

Publications and source records attributed to G Rizzolatti.

At least 55 records · Page 3Linked to original sources

Convergence of pallidal and cerebellar outputs on the frontal motor areas.

The general pattern of the organization of the thalamo-cortical projections to the agranular frontal cortex of the monkey is still matter of debate. An important issue is whether each motor area is the target of a single thalamic nucleus or it receives afferences from multiple thalamic sources. In this light it is of interest to determine whether the basal ganglia and cerebellar outputs, which are segregated in the thalamus, remain segregated also at the cortical level or, on the contrary, both converge on the same cortical areas. In the present article we present data concerning the thalamic input to mesial area 6 obtained with cortical injections of retrograde neural tracers. This cortical sector, classically considered as coextensive with the so called supplementary motor area (SMA), was recently found to be formed by two independent anatomo-functional areas: F3 (SMA-proper) and F6 (pre-SMA). On the basis of the neurophysiological properties of the two areas we have proposed that F6 plays a hierarchically higher role in motor control than F3. The present results allow us the following main conclusions: a) Each motor area is the target of a distinct set of thalamic nuclei. b) Each area is the target of both basal ganglia and cerebellar outflows. c) As far as the basal ganglia input is concerned, F3 is a part of the so called "basal ganglia motor loop", whereas, F6 belongs to the "basal ganglia complex loop". This differential basal ganglia input provides further evidence in favor of a higher hierarchical role of F6 in comparison to F3.

Animals↗

Corticospinal projections from mesial frontal and cingulate areas in the monkey.

We injected neural tracers into the lateral funiculus of the spinal cord in order to relate the sites of origin of the spinal projections from the mesial cortical surface with the cytoarchitectonic organization of this region. We found a close correlation between the origin sites and density of corticospinal projections and the areal organization. The areas most densely labelled were F3 (SMA-proper) and area 24d, whereas F6 (pre-SMA) and area 24c showed a low density of labelling. The segmental topography of the corticospinal projections fitted well with the somatotopy of the mesial cortical areas. We conclude that in the agranular mesial cortex there are four independent motor representations: F3 and 24d where the whole body is represented, and F6 and 24c which are mostly related to arm movements.

Amidines↗

The fronto-parietal cortex of the prosimian Galago: patterns of cytochrome oxidase activity and motor maps.

We mapped the motor areas of the prosimian Galago crassicaudatus using intracortical electrical microstimulation and morphological and histochemical (cytochrome oxidase) techniques. Stimulation data showed that on the brain convexity there is an area (area Frontalis posterior, F post.) from which movements could be evoked at low threshold (< 10 microA). This area is somatotopically organized, with the leg represented medially, the arm centrally and the face and mouth laterally. Proximal and distal movements are not segregated. Most of the evoked movements, even at threshold, consist of movements involving two or more joints. F post. is characterized by a three-band cytochrome oxidase activity pattern. It has an agranular structure, but it lacks pyramidal cells that are larger than those observed in other areas. In front of F post. there is an area histochemically similar to it, Frontalis intermedialis (F int.). This area consists of two cytoarchitectonic divisions: an agranular division (F int. pars caudalis) and a disgranular division (F int. pars rostralis). The excitability threshold of F int. is relatively high (10 to 30 microA). Eye, ear and neck movements are elicited from its lateral part, whereas trunk movements associated with limb movements are elicited from its medial part. Caudal to F post., there is another region from which movements can be evoked with currents between 10 to 30 microA. This region has the same medio-lateral somatotopic arrangement of F post. Typically, single joint movements are elicited from it. Proximal and distal movements are not segregated. In spite of its homogeneity in terms of motor response, the posterior excitable region is formed by two anatomically separate areas: anterior somatic area (S ant.) and posterior somatic area (S post.). S ant. has a typical koniocortex structure, whereas S post, resembles the parakoniocortex as defined by Sanides (J. Hirnforsch., 9 (1967) 225-252). Histochemically both areas are made up of four longitudinal stripes differing for enzymatic activity. The three superficial stripes tend to merge together and are sharply separated from a deeply located, light stripe. This stripe is homogeneous in S ant., whilst its central part shows an increase in activity in S post. The possible homologies between the motor and somatic areas of the galago and monkey as well as their role in movement control are discussed.

Animals↗

Orienting of attention and eye movements.

According to the premotor theory of attention, the mechanisms responsible for spatial attention and the mechanisms involved in programming ocular saccades are basically the same. The aim of the present experiments was to test this claim. In experiment 1 subjects were presented with a visual display consisting of a fixation point and four boxes arranged horizontally and located above the fixation cross. Two of the boxes were in the left visual hemifield, two in the right. A fifth box was located on the vertical meridian below the fixation cross. Digit cues indicated in which of the upper boxes the imperative stimulus was most likely to appear. Subjects were instructed to direct attention to the cued box and to perform a saccadic eye movement to the lower box on presentation of the imperative stimulus. The trajectory of the saccades deviated contralateral to the hemifield in which the imperative stimulus was presented. This deviation was larger when the hemifield where the imperative stimulus was presented was the cued one. In experiment 2, the visual display consisted of five boxes forming a cross. The central box served as a fixation point. The cue was a small line, linked to the central box, pointing to different directions and indicating where the visual imperative stimulus would appear. In 50% of trials, the imperative stimulus was a visual stimulus presented either in one of the lateral boxes or in the central one. In the remaining 50% of trials, the imperative stimulus was a non-lateralised sound. Half the subjects were instructed to make a saccade to the upper box at the presentation of the visual imperative stimulus and to the lower box at the presentation of the acoustic stimulus. Half the subjects received the opposite instructions. The results confirmed that the saccades deviate contralateral to the hemifield of stimulus presentation in the case of visual imperative stimuli. Most importantly, they showed that the saccades deviate contralateral to the cued hemifield, also in the case of acoustic imperative stimuli. Experiment 3 was similar to experiment 2. It confirmed the results of that experiment and showed that slow ocular drifts, which are observed in the time interval between cue and imperative stimulus presentation, cannot explain the ocular deviations. Taken together, the experiments demonstrate that spatial attention allocation leads to an activation of oculomotor circuits, in spite of eye immobility.

Attention↗

Corticocortical connections of area F3 (SMA-proper) and area F6 (pre-SMA) in the macaque monkey.

The monkey mesial area 6 comprises two distinct cytoarchitectonic areas: F3 [supplementary motor area properly defined (SMA-proper)], located caudally, and F6 (pre-SMA), located rostrally. The aim of the present study was to describe the corticocortical connections of these two areas. To this purpose restricted injections of neuronal tracers (wheat germ-agglutinin conjugated to horseradish peroxidase, fluorescent tracers) were made in different somatotopic fields of F3, F6, and F1 (area 4) and their transport plotted. The results showed that F3 and F6 differ markedly in their cortical connections. F3 is richly linked with F1 and the posterior premotor and cingulate areas (F2, F4, 24d). Connections with the anterior premotor and cingulate areas (F6, F7, F5, 24c) although present, are relatively modest. There is no input from the prefrontal lobe. F3 is also connected with several postrolandic cortical areas. These connections are with areas PC, PE, and PEa in the superior parietal lobule, cingulate areas 23 and PEci, the opercular parietal areas (PFop, PGop, SII) and the granular insula. F6 receives a rich input from the anterior premotor areas (especially F5) and cingulate area 24c, whereas its input from the posterior premotor and cingulate areas is very weak. A strong input originates from area 46. There are no connections with F1. The connections with the postrolandic areas are extremely meagre. They are with areas PG and PFG in the inferior parietal lobule, the disgranular insula, and the superior temporal sulcus. A further result was the demonstration of a differential connectivity pattern of the cingulate areas 24d and 24c. Area 24d is strongly linked with F1 and F3, whereas area 24c is connected mostly with F6. The present data support the notion that the classical SMA comprises two functionally distinct areas. They suggest that F6 (the rostral area) is responsible for the "SMA" so-called high level motor functions, whereas F3 (the caudal area) is more closely related to movement execution.

Animals↗

Activation of precentral and mesial motor areas during the execution of elementary proximal and distal arm movements: a PET study.

Regional cerebral blood flow was measured using positron emission tomography (PET) in normal subjects while performing simple aimless proximal and distal arm movements. The aim of the experiment was to compare the somatotopic organization of precentral and mesial (the so called supplementary motor area, SMA) motor cortices and to evaluate whether in man, as in the monkey, the rostral and caudal sectors of SMA are functionally different. The results showed that proximal and distal arm movements are to a large extent segregated in the precentral motor cortex, but not in the SMA. They also showed that the SMA is made of at least two functional sectors. Only the caudal one is activated during simple aimless movements.

Adult↗

Study of selective reaching and grasping in a patient with unilateral parietal lesion. Dissociated effects of residual spatial neglect.

In the present study we investigated the possibility of a dissociation between the visual control of reaching and the visual control of grasping in a prehension task. To this purpose we studied the kinematics of prehension movements in a patient with a right parietal lesion and in six right-handed healthy control subjects. The task we used was one in which the subjects had to reach and grasp target objects in the presence or absence of a simultaneously presented distractor object. All stimuli were presented in the space ipsilateral to the lesion. The distractor could be either of the same or different size to the target object and was presented either to the right or to the left of the target. The following parameters of the prehension 'transport' component were analysed: wrist trajectory, transport time, tangential peak velocity, acceleration. Maximal finger aperture, time to maximal finger aperture, peak acceleration and time to peak acceleration of grip aperture were the parameters of the 'grasping' component analysed. The results showed that, although the patient had no misreaching, her hand trajectory deviated abnormally towards the distractor position when the distractor was to the right (ipsilateral) side of the target. In contrast, the grasp kinematics was not affected by the distractors, even when the size of the right distractor was different from the target. It appears, therefore, that the attentional shift towards the ipsilesional side, typical of neglect patients, determines a surprising dissociation in motor control. In the presence of a right distractor, the patient plans and partially executes a reaching movement towards that object and simultaneously performs a grasping movement towards a second object, i.e. the centrally located target. The presentation of distractors had no effects on the prehension kinematics of the control subjects.

Attention↗

European isolation and confinement study. Attention during isolation and confinement.

The aim of this study was to examine the effect of isolation and confinement on selective attention. The experiments were carried out on six specially selected male subjects. The subjects were isolated in a hyperbaric chamber complex at a low overpressure for a period of 28 days. Distribution of attention between two tasks, visual search and orientation of attention to symbolic and automatic cues were studied. The results showed that 4 weeks of isolation and confinement do not significantly modify visual search and the capacity to execute two tasks simultaneously. In the double task experiment there was a clear improvement of performance during isolation and the post-isolation period. However, the development of a remarkably large difference between subjects was found. Orientation of attention to automatic cues gave results comparable to those obtained under normal conditions. Valid trials (stimulus flash and cue in same box) were faster than both neutral (cue in all boxes) and invalid trials (stimulus flash and cue in different boxes). Orientation of attention to symbolic cues also produced this effect. However, the "meridian effect" (increased reaction time when the stimulus is in the opposite half of the visual field as cue), which is constantly observed under normal conditions, was absent during isolation. The absence of the meridian effect is interpreted as a change in the subject's strategy to allocate attention. It is proposed that the absence of the meridian effect indicates an initial deficit of attentional capacities during isolation and confinement.

Adult↗

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↗