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A P Georgopoulos

Publications and source records attributed to A P Georgopoulos.

At least 19 recordsLinked to original sources

Motor cortical activity in a context-recall task.

A monkey was trained to respond on the basis of the serial position of a test stimulus in a sequence. First, three stimuli were presented successively on a circle. Then one of them (except the last) changed color (test stimulus) and served as the go signal: The monkey was required to produce a motor response in the direction of the stimulus that followed the test stimulus. When the test stimulus was the second in the sequence, there was a change in motor cortical activity from a pattern reflecting the direction of this stimulus to the pattern associated with the direction of the motor response. This change was abrupt, occurred 100 to 150 milliseconds after the go signal, and was evident both in the activity of single cells and in the time-varying neuronal population vector. These findings identify the neural correlates of a switching process that is different from a mental rotation described previously.

Animals

Overlapping neural networks for multiple motor engrams.

The hypothesis was tested that learned movement trajectories of different shapes can be stored in, and generated by, largely overlapping neural networks. Indeed, it was possible to train a massively interconnected neural network to generate different shapes of internally stored, dynamically evolving movement trajectories using a general-purpose core part, common to all networks, and a special-purpose part, specific for a particular trajectory. The weights of connections between the core units do not carry any information about trajectories. The core network alone could generate externally instructed trajectories but not internally stored ones, for which both the core and the trajectory-specific part were needed. All information about the movements is stored in the weights of connections between the core part and the specialized units and between the specialized units themselves. Due to these connections the core part reveals specific dynamical behavior for a particular trajectory and, as the result, discriminates different tasks. The percentage of trajectory-specific units needed to generate a certain trajectory was small (2-5%), and the total output of the network is almost entirely provided by the core part, whereas the role of the small specialized parts is to drive the dynamical behavior. These results suggest an efficient and effective mechanism for storing learned motor patterns in, and reproducing them by, overlapping neural networks and are in accord with neurophysiological findings of trajectory-specific cells and with neurological observations of loss of specific motor skills in the presence of otherwise intact motor control.

Models, Theoretical

Directional operations in the motor cortex modeled by a neural network of spiking neurons.

A neural network with realistically modeled, spiking neurons is proposed to model ensemble operations of directionally tuned neurons in the motor cortex. The model reproduces well directional operations previously identified experimentally, including the prediction of the direction of an upcoming movement in reaching tasks and the rotation of the neuronal population vector in a directional transformation task.

Action Potentials

Behavioral neurophysiology of the motor cortex.

The study of the motor cortex in behaving monkeys during the past 20 years has provided important information on the brain mechanisms underlying motor control. With respect to reaching movements in space, several aspects of motor cortical function concerning the specification of the direction of movement have now been elucidated and are reviewed in this article. The activity of single cells in the motor cortex is broadly tuned with respect to the direction of reaching, so that the discharge rate is highest with movements in a preferred direction and decreases progressively with movements made in directions more and more away from the preferred one. Thus the neural command for the direction of reaching can be regarded as an ensemble of cell vectors, with each vector pointing in the cell's preferred direction and having a length proportional to the change in cell activity. The outcome of this population code can be visualized as a vector that points in the direction of the upcoming movement during the reaction time, during an instructed delay period, and during a memorized delay period. Moreover, when a mental transformation is required for the generation of a reaching movement in a different direction from a reference direction, the population vector provides a direct insight into the nature of the cognitive process by which the required transformation is achieved.

Animals

The motor cortex and the coding of force.

The relation of cellular activity in the motor cortex to the direction of two-dimensional isometric force was investigated under dynamic conditions in monkeys. A task was designed so that three force variables were dissociated: the force exerted by the subject, the net force, and the change in force. Recordings of neuronal activity in the motor cortex revealed that the activity of single cells was directionally tuned and that this tuning was invariant across different directions of a bias force. Cell activity was not related to the direction of force exerted by the subject, which changed drastically as the bias force changed. In contrast, the direction of net force, the direction of force change, and the visually instructed direction all remained quite invariant and congruent and could be the directional variables, alone or in combination, to which cell activity might relate.

Animals

Three-dimensional drawings in isometric conditions: planar segmentation of force trajectory.

Normal human subjects grasped an isometric handle with an unrestrained, pronated hand. They were asked to exert forces continuously to draw lemniscates (figure eights) in specified or self-chosen planes and in the presence or absence of a three-dimensional visual feedback cursor and a visual template. In every condition, the mean plane orientation in the force space differed appreciably between the two loops of the figure, as described previously by Soechting and Terzuolo (1987a) for free drawing arm movements. These findings suggest that the planar segmentation of the motor trajectory is not a consequence of joint motion but arises from central constraints related to the production of motor trajectory in space.

Feedback

Three-dimensional drawings in isometric conditions: relation between geometry and kinematics.

Normal human subjects grasped a 3-D isometric handle with an otherwise unrestrained, pronated hand and exerted forces continuously to draw circles, ellipses and lemniscates (figure-eights) in specified planes in the presence or absence of a 3-D visual force-feedback cursor and a visual template. Under any of these conditions and in all subjects, a significant positive correlation was observed between the instantaneous curvature and angular velocity, and between the instantaneous radius of curvature and tangential velocity; that is, when the force trajectory was most curved, the tangential velocity was lowest. This finding is similar to that obtained by Viviani and Terzuolo (1982) for 2-D drawing arm movements and supports the notion that central constraints give rise to the relation between geometric and kinematic parameters of the trajectory.

Female

Motor cortical activity in a memorized delay task.

Two rhesus monkeys were trained to move a handle on a two-dimensional (2D) working surface in directions specified by a light at the plane. They first captured with the handle a light on the center of the plane and then moved the handle in the direction indicated by a peripheral light (cue signal). The signal to move (go signal) was given by turning off the center light. The following tasks were used: (a) In the non-delay task the peripheral light was turned on at the same time as the center light went off. (b) In the memorized delay task the peripheral light stayed on for 300 ms and the center light was turned off 450-750 ms later. Finally, (c) in the non-memorized delay task the peripheral light stayed on continuously whereas the center light went off 750-1050 ms after the peripheral light came on. Recordings in the arm area of the motor cortex (N = 171 cells) showed changes in single cell activity in all tasks. In both delay tasks, the neuronal population vector calculated every 20 ms after the onset of the peripheral light pointed in the direction of the upcoming movement, which was instructed by the cue light. Moreover, the strength of the population signal showed an initial peak shortly after the cue onset in both the memorized and non-memorized delay tasks but it maintained a higher level during the memorized delay period, as compared to the non-memorized task.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals

Cognitive spatial-motor processes. 7. The making of movements at an angle from a stimulus direction: studies of motor cortical activity at the single cell and population levels.

Two rhesus monkeys were trained to move a handle on a two-dimensional (2-D) working surface either towards a visual stimulus ("direct" task) or in a direction orthogonal and counterclockwise (CCW) from the stimulus ("transformation" task), depending on whether the stimulus appeared dim or bright, respectively. Thus the direction of the stimulus (S, in polar coordinates) and the direction of the movement (M) were the same in the direct task but differed in the transformation task, such that M = S + 90 degrees CCW. The task (i.e. brightness) condition (k = 2, i.e. direct or transformation) and the direction of the stimulus (m = 8, i.e. 8 equally spaced directions on a circle) resulted in 16 combinations (k x m = 16 "classes") that were varied from trial to trial in a randomized block design. In 8 of these combinations the direction of the stimulus was the same for both tasks, whereas the direction of the movement was the same in the remaining 8 cases. The electrical signs of cell activity (N = 394 cells) in the arm area of the motor cortex (contralateral to the performing arm) were recorded extracellularly. The neural activity was analyzed at the single cell and neuronal population levels, and a modeling of the time course of single activity during the transformation task was carried out. We found the following. (a) Individual cells were active in both tasks; no cells were found that were active exclusively in only one of the two tasks. The patterns of single cell activity in the transformation task frequently differed from those observed in the direct task when the stimulus or the movement were the same. More specifically, cells could not be consistently classified as "movement"-or "stimulus"-related for frequently the activity of a particular cell would seem "movement-related" for a particular stimulus-movement combination, "stimulus-related" for another combination, or unrelated to either movement or stimulus for still another combination. Thus no real insight could be gained from such an analysis of single cell activity. (e) In a different analysis, we explored the idea that a changing directional signal could be detected in the time course of single cell activity during the reaction time. For that purpose we modeled the time course of single activity observed in the transformation task as a linear, weighted combination of influences from the direct task, taking the time patterns of cell activity during the stimulus, intermediate and movement directions in the direct task as estimates of the postulated directional influences.(ABSTRACT TRUNCATED AT 400 WORDS)

Analysis of Variance

Cognitive spatial-motor processes. 4. Specification of the direction of visually guided isometric forces in two-dimensional space: information transmitted and effects of visual force-feedback.

The information transmitted (Ti) by the direction of two-dimensional (2-D) isometric forces at different stereoscopic depths was studied in 50 naive human subjects using an isometric manipulandum and random dot stereograms generated in a color display (Massey et al. 1988). Subjects viewed the display through appropriate color filters and perceived the image of a disk rotated about a horizontal axis on the frontal plane; the top of the disk was rotated around that axis by 15, 45, 60 and 80 degrees away from the subject. Each of these disks involved a different amount of stereoscopic depth perception which was lowest for the 15 degrees and highest for the 80 degrees tilt. Subjects were instructed to exert force in the direction of a visual target presented on the disk in a reaction time task. The instantaneous force exerted by the subjects on the manipulandum was shown on the disk in the form of a feedback cursor. Information transmitted, reaction time (RT) and systematic directional deviations were calculated. We found the following. (a) Ti increased with input information but at a lower rate; at the highest level of input information studied (5.91 bits), Ti was 4.1 bits at the 15 degrees tilt. This high value of Ti suggests that directional information for isometric force is processed very efficiently. However, this Ti was consistently lower than that transmitted by the direction of movement (Georgopoulos and Massay, 1988). (b) Ti did not differ significantly among the 15-60 degrees tilt but was 0.19 bits less for the 80 degrees tilt. RT did not differ among the 15-80 degrees tilts.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult

Cognitive spatial-motor processes. 5. Specification of the direction of visually guided isometric forces in two-dimensional space: time course of information transmitted and effect of constant force bias.

The effects of an external constant force bias on the information transmitted (Ti) by the direction of isometric force exerted in 2-dimensional (2-D) space by human subjects were studied using an isometric manipulandum and random dot stereograms generated in a color display (Massey et al. 1988, Massey et al. 1990). Subjects exerted force on the manipulandum such that a visual force-feedback cursor would move in the direction of a visually defined stimulus in the stereo display. The time course of force development and the gain of directional information during increasing force intensity were also studied. We found the following. (a) When no bias force was applied, the force exerted by the subject increased from near zero to greater than 200 gram-force at the end of a trial and was close to the visually defined direction. When a constant bias force of 110 gram-force was applied in various directions in blocks of trials, the force exerted by the subject increased in time, as above; however, its direction also changed in time so that the instantaneous vector sum of the bias force and the force exerted by the subject pointed close to the visually defined direction. The Ti and the reaction time (RT) did not differ significantly in the two experimental conditions. These results suggest that the directional control of isometric forces is very efficient, especially in relation to visuomotor coordination. (b) The Ti was calculated at various levels of force intensity, as the latter increased from approximately 50 gram-force to 200 gram-force.(ABSTRACT TRUNCATED AT 250 WORDS)

Cognition

Cognitive spatial-motor processes. 6. Visuomotor memory scanning.

Fourteen human subjects performed in a modified Sternberg memory-scanning task. First, they made a series of 2-6 movements in different directions from a central point towards peripheral lights on a planar working surface ("list trials"). Then, after a warning signal, one of the previous list stimuli, except the last, was presented again ("test trial"). Subjects were instructed to move in the direction of the stimulus which was presented next in sequence in the list. The mean reaction time (RT) in the test trials increased as a linear function of the number of movements, S, in the list: Mean RT (ms) = 105 + 205.8S (2 less than or equal to S less than or equal to 6). This finding suggests that the task involves memory scanning of visuomotor list items.

Cognition

Cortical control of motor behavior at the cellular level.

The studies reviewed in this paper describe the relations of single-cell activity in central motor structures to complex visuomotor tasks and document the fact that various cortical areas process visuomotor information in parallel. Moreover, the studies provide clear evidence that the map in the motor cortex is modifiable and dynamically maintained.

Animals

Parietal cortex neurons of the monkey related to the visual guidance of hand movement.

A class of neurons specifically related to hand movements was studied in the posterior parietal cortex while the monkeys manipulated different types of objects. We examined the neuronal activity during manipulation of objects by the hand in the light and in the dark. Fifty-five neurons were active during manipulation in the dark and were classified as "hand-movement-related" neurons. Of these, 38/55 (69%) cells were also influenced by the visual stimulus. Most of the hand-movement-related neurons were selective in the type of objects manipulated. Moreover, some of these cells were selective in the axis of orientation of the object. These results suggest that the hand-movement-related neurons of the parietal cortex are concerned with the visual guidance of the hand movement, especially in matching the pattern of movement with the spatial characteristics of the object to be manipulated.

Animals