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Biomedical subjects

G Rizzolatti

Publications and source records attributed to G Rizzolatti.

At least 73 records · Page 4Linked to original sources

[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↗

Functional organization of inferior area 6 in the macaque monkey. II. Area F5 and the control of distal movements.

The functional properties of neurons located in the rostral part of inferior area 6 were studied in awake, partially restrained macaque monkeys. The most interesting property of these neurons was that their firing correlated with specific goal-related motor acts rather than with single movements made by the animal. Using the motor acts as the classification criterion we subdivided the neurons into six classes, four related to distal motor acts and two related to proximal motor acts. The distal classes are: "Grasping-with-the-hand-and-the-mouth neurons", "Grasping-with-the-hand neurons", "Holding neurons" and "Tearing neurons". The proximal classes are: "Reaching neurons" and "Bringing-to-the-mouth-or-to-the-body neurons". The vast majority of the cells belonged to the distal classes. A particularly interesting aspect of distal class neurons was that the discharge of many of them depended on the way in which the hand was shaped during the motor act. Three main groups of neurons were distinguished: "Precision grip neurons", "Finger prehension neurons", "Whole hand prehension neurons". Almost the totality of neurons fired during motor acts performed with either hand. About 50% of the recorded neurons responded to somatosensory stimuli and about 20% to visual stimuli. Visual neurons were more difficult to trigger than the corresponding neurons located in the caudal part of inferior area 6 (area F4). They required motivationally meaningful stimuli and for some of them the size of the stimulus was also critical. In the case of distal neurons there was a relationship between the type of prehension coded by the cells and the size of the stimulus effective in triggering the neurons. It is proposed that the different classes of neurons form a vocabulary of motor acts and that this vocabulary can be assessed by somatosensory and visual stimuli.

Animals↗

Right hemisphere superiority for programming oculomotion: evidence from simple reaction time experiments.

Simple reaction times (RTs) to lateralized unstructured visual stimuli were measured in normal subjects while they were carrying out concomitant oculomotor tasks. Four tasks were used. In the first task, subjects had to find the correct path in a maze presented at the centre of a screen; in the second task, subjects had to follow a bright dot moved on a screen with variable direction, trajectory and velocity with their eyes; in the third task, the subjects had to follow a bright dot moved back and forth either horizontally or vertically along the same trajectory and at a constant velocity; in the fourth task, the subjects had to monitor the movement of a bright dot moved with variable direction, trajectory and velocity without moving the eyes. In all tasks, with the exception of the third, there was a selective lengthening of RTs mediated by the right hemisphere. It is concluded that the right hemisphere is dominant in programming eye movements.

Adult↗

Functional organization of inferior area 6.

The rostral part of the agranular frontal cortex (area 6) of the monkey consists of two large sectors: a superior sector lying medial to the spur of the arcuate sulcus (superior area 6) and an inferior sector lying lateral to it (inferior area 6). Single neurons have been recorded from inferior area 6 in behaving monkeys (Macaca nemestrina). The results were: (a) Proximal movements are essentially represented caudally in the histochemically defined area F4. Neurons related to these movements respond strongly to tactile and visual stimuli. Visual receptive fields are located in the space around the animal's body (peripersonal space) and their location does not change with eye movements. The direction of movements effective in triggering the neurons is congruent with the position of their visual receptive field. (b) Distal movements are represented rostrally in the anterior part of F4 and in F5. Neurons related to these movements discharge vigorously during motor acts that have a precise aim. The neurons were subdivided into four classes: grasping-with-the-hand neurons, grasping-with-the-hand-and-mouth neurons, holding neurons, and tearing neurons. Regardless of the class they belong to a large number of neurons show specificity for different types of object prehension--discharging, for example, during precision grip but not during whole-hand prehension. It is proposed that inferior area 6 contains a vocabulary of motor acts related to hand-mouth movements. The motor acts can be retrieved by visual and somatosensory stimuli. The possibility is discussed that a series of vocabularies where movements of various complexity are stored represents the neural basis of cortical motor organization.

Animals↗

Neurons related to goal-directed motor acts in inferior area 6 of the macaque monkey.

A new class of neurons was identified in the rostral part of inferior area 6 in the macaque monkey (Macaca nemestrina). These neurons fire in relation to motor acts which have a particular aim such as reaching, grasping or holding. The same neuron discharges when the animal uses the right hand, the left hand and, frequently, also the mouth. Furthermore most of these neurons specify how the aim can be achieved (e.g. precision grip vs whole hand prehension). Different types of goal-related neurons form a vocabulary of simple motor acts localized in inferior area 6.

Action Potentials↗

Movements of attention in the three spatial dimensions and the meaning of "neutral" cues.

Six experiments were conducted to examine the effect of various attentional manipulations on reaction time to visual stimuli. The first three experiments compared the responses to stimuli presented in the depth (Experiment 1), along the horizontal (Experiment 2), and vertical (Experiment 3) meridians in a valid condition (stimulus presented in the cued position), an invalid condition (stimulus presented in the alternative position to the cued position) and a neutral condition (no information on stimulus position). The most interesting result was the demonstration that attention can be moved along the sagittal plane in the absence of vergence eye movements and that when attention is focused on a certain point, unattended points between this point and the observer (i.e. near points) are responded faster than unattended points beyond it (i.e. far points). In the frontal plane no asymmetry was found between the responses to unattended points above or below the fixation, whereas a certain, albeit non-constant, advantage was present for unattended stimuli on the right of the fixation point in respect to those on the left of it. The second series of experiments was similar to the first one, except that a new situation was introduced in which the fixation point was cued and stimuli could appear either in correspondence to it or in a peripheral position (invalid condition with attention at the fixation point). The results showed that in this new situation the responses to unattended stimuli are much longer than they are under neutral conditions, and as long as they are under conventional invalid condition. It is suggested that the so called neutral condition is a condition of diffuse attention and an attempt is made to explain it in terms of a premotor theory of attention.

Attention↗

Reorienting attention across the horizontal and vertical meridians: evidence in favor of a premotor theory of attention.

Stimuli presented in a non-attended location are responded to much slower than stimuli presented in an attended one. The hypotheses proposed to explain this effect make reference to covert movement of attention, hemifield inhibition, or attentional gradients. The experiment reported here was aimed at discriminating among these hypotheses. Subjects were cued to attend to one of four possible stimulus locations, which were arranged either horizontally or vertically, above, below, to the right or left of a fixation point. The instructions were to respond manually as fast as possible to the occurrence of a visual stimulus, regardless of whether it occurred in a cued or in a non-cued location. In 70% of the cued trials the stimulus was presented in the cued location and in 30% in one of the non-cued locations. In addition there were trials in which a non-directional cue instructed the subject to pay attention to all four locations. The results showed that the correct orienting of attention yielded a small but significant benefit; the incorrect orienting of attention yielded a large and significant cost; the cost tended to increase as a function of the distance between the attended location and the location that was actually stimulated; and an additional cost was incurred when the stimulated and attended locations were on opposite sides of the vertical or horizontal meridian. We concluded that neither the hypothesis postulating hemifield inhibition nor that postulating movement of attention with a constant time can explain the data. The hypothesis of an attention gradient and that of attention movements with a constant speed are tenable in principle, but they fail to account for the effect of crossing the horizontal and vertical meridians. A hypothesis is proposed that postulates a strict link between covert orienting of attention and programming explicit ocular movements. Attention is oriented to a given point when the oculomotor programme for moving the eyes to this point is ready to be executed. Attentional cost is the time required to erase one ocular program and prepare the next one.

Attention↗