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John F Kalaska

Publications and source records attributed to John F Kalaska.

11 recordsLinked to original sources

Sequential activation of muscle synergies during locomotion in the intact cat as revealed by cluster analysis and direct decomposition.

During goal-directed locomotion, descending signals from supraspinal structures act through spinal interneuron pathways to effect modifications of muscle activity that are appropriate to the task requirements. Recent studies using decomposition methods suggest that this control might be facilitated by activating synergies organized at the level of the spinal cord. However, it is difficult to directly relate these mathematically defined synergies to the patterns of electromyographic activity observed in the original recordings. To address this issue, we have used a novel cluster analysis to make a detailed study of the organization of the synergistic patterns of muscle activity observed in the fore- and hindlimb during treadmill locomotion. The results show that the activity of a large number of forelimb muscles (26 bursts of activity from 18 muscles) can be grouped into 11 clusters on the basis of synchronous co-activation. Nine (9/11) of these clusters defined muscle activity during the swing phase of locomotion; these clusters were distributed in a sequential manner and were related to discrete behavioral events. A comparison with the synergies identified by linear decomposition methods showed some striking similarities between the synergies identified by the different methods. In the hindlimb, a simpler organization was observed, and a sequential activation of muscles similar to that observed in the forelimb during swing was less clear. We suggest that this organization of synergistic muscles provides a means by which descending signals could provide the detailed control of different muscle groups that is necessary for the flexible control of multi-articular movements.

Animals↗

Parietal area 5 activity does not reflect the differential time-course of motor output kinetics during arm-reaching and isometric-force tasks.

Many single-neuron recording studies have examined the degree to which the activity of primary motor cortex (M1) neurons is related to the kinematics and kinetics of various motor tasks. This has not been explored as extensively for arm movement-related neurons in posterior parietal cortex area 5. We recorded the activity of 78 proximal arm-related neurons in area 5 of two monkeys while they used their whole arm to make reaching movements toward eight targets on a horizontal plane against an inertial load or to generate isometric forces at the hand in the same eight horizontal directions. The overall range of measured output forces was similar in the two tasks. The forces increased monotonically in the desired direction in the isometric task. In the movement task, in contrast, they showed a rapid initial increase in the direction of movement, followed by a transient reversal of forces as the hand approached the target. Many task-related area 5 neurons were tuned for the direction of motor output in the tasks, but most area 5 neurons were more strongly active or exclusively active in the movement task than in the isometric task. Furthermore, their activity at either the single cell or population level did not reflect the transient reversal of output forces during movement. In contrast, M1 neuronal activity was typically strong in both tasks and showed task-related changes that reflected the differences in the time course and directionality of force outputs between both tasks, including the transient reversal of forces in the movement task. These results show that area 5 neurons are less strongly related to the time-course of task kinetics than M1 during isometric and arm-movement tasks.

Action Potentials↗

Motor cortex neural correlates of output kinematics and kinetics during isometric-force and arm-reaching tasks.

We recorded the activity of 132 proximal-arm-related neurons in caudal primary motor cortex (M1) of two monkeys while they generated either isometric forces against a rigid handle or arm movements with a heavy movable handle, in the same eight directions in a horizontal plane. The isometric forces increased in monotonic fashion in the direction of the force target. The forces exerted against the handle in the movement task were more complex, including an initial accelerating force in the direction of movement followed by a transient decelerating force opposite to the direction of movement as the hand approached the target. EMG activity of proximal-arm muscles reflected the difference in task dynamics, showing directional ramplike activity changes in the isometric task and reciprocally tuned "triphasic" patterns in the movement task. The apparent instantaneous directionality of muscle activity, when expressed in hand-centered spatial coordinates, remained relatively stable during the isometric ramps but often showed a large transient shift during deceleration of the arm movements. Single-neuron and population-level activity in M1 showed similar task-dependent changes in temporal pattern and instantaneous directionality. The momentary dissociation of the directionality of neuronal discharge and movement kinematics during deceleration indicated that the activity of many arm-related M1 neurons is not coupled only to the direction and speed of hand motion. These results also demonstrate that population-level signals reflecting the dynamics of motor tasks and of interactions with objects in the environment are available in caudal M1. This task-dynamics signal could greatly enhance the performance capabilities of neuroprosthetic controllers.

Animals↗

Neural correlates of reaching decisions in dorsal premotor cortex: specification of multiple direction choices and final selection of action.

We show that while a primate chooses between two reaching actions, its motor system first represents both options and later reflects selection between them. When two potential targets appeared, many (43%) task-related, directionally tuned cells in dorsal premotor cortex (PMd) discharged if one of the targets was near their preferred direction. At the population level, this generated two simultaneous sustained directional signals corresponding to the current reach options. After a subsequent nonspatial cue identified the correct target, the corresponding directional signal increased, and the signal for the rejected target was suppressed. The PMd population reliably predicted the monkey's response choice, including errors. This supports a planning model in which multiple reach options are initially specified and then gradually eliminated in a competition for overt execution, as more information accumulates.

Animals↗

Neural correlates of mental rehearsal in dorsal premotor cortex.

Behavioural and imaging studies suggest that when humans mentally rehearse a familiar action they execute some of the same neural operations used during overt motor performance. Similarly, neural activation is present during action observation in many of the same brain regions normally used for performance, including premotor cortex. Here we present behavioural evidence that monkeys also engage in mental rehearsal during the observation of sensory events associated with a well-learned motor task. Furthermore, most task-related neurons in dorsal premotor cortex exhibit the same activity patterns during observation as during performance, even during an instructed-delay period before any actual observed motion. This activity might be a single-neuron correlate of covert mental rehearsal.

Animals↗

Systematic changes in motor cortex cell activity with arm posture during directional isometric force generation.

We report here the activity of 96 cells in primate primary motor cortex (MI) during exertion of isometric forces at the hand in constant spatial directions, while the hand was at five to nine different spatial locations on a plane. The discharge of nearly all cells varied significantly with both hand location and the direction of isometric force before and during force-ramp generation as well as during static force-hold. In addition, nearly all cells displayed changes in the variation of their activity with force direction at different hand locations. This change in relationship was often expressed in part as a change in the cell's directional tuning at different hand locations. Cell directional tuning tended to shift systematically with hand location even though the direction of static force output at the hand remained constant. These directional effects were less pronounced before the onset of force output than after force onset. Cells also often showed planar modulations of discharge level with hand location. Sixteen proximal arm muscles showed similar effects, reflecting how hand location-dependent biomechanical factors altered their task-related activity. These findings indicate that MI single-cell activity does not covary exclusively with the level and direction of net force output at the hand and provides further evidence that MI contributes to the transformation between extrinsic and intrinsic representations of motor output during isometric force production.

Animals↗

Neural activity in primary motor and dorsal premotor cortex in reaching tasks with the contralateral versus ipsilateral arm.

To investigate the effector dependence of task-related neural activity in dorsal premotor (PMd) and primary motor cortex (M1), directional tuning functions were compared between instructed-delay reaching tasks performed separately with either the contralateral or the ipsilateral limb. During presentation of the instructional cue, the majority (55/90, 61%) of cells in PMd were tuned with both arms, and their dynamic range showed a trend for stronger discharge with the contralateral arm. Most strikingly, however, the preferred direction of most of these latter cells (41/55, 75%) was not significantly different between arms. During movement, many PMd cells continued to be tuned with both arms (53/90, 59%), with a trend for increasing directional differences between the arms over the course of the trial. In contrast, during presentation of the instructional cue only 5/74 (7%) cells in M1 were tuned with both arms. During movement, about half of M1 cells (41/74, 55%) were tuned with both arms but the preferred directions of their tuning functions were often very different and there was a strong bias toward greater discharge rates when the contralateral arm was used. Similar trends were observed for EMG activity. In conclusion, M1 is strongly activated during movements of the contralateral arm, but activity during ipsilateral arm movements is also common and usually different from that seen with the contralateral arm. In contrast, a major component of task-related activity in PMd represents movement in a more abstract or task-dependent and effector-independent manner, especially during the instructed-delay period.

Animals↗

Context-dependent anticipation of different task dynamics: rapid recall of appropriate motor skills using visual cues.

Recent studies have reported that human subjects show varying degrees of ability to use contextual cues to recall the motor skills required to compensate for different dynamic external force fields during arm movements. In particular, the subjects showed little or no ability to use color cues to adjust motor outputs in anticipation of the perturbing fields after limited periods of training that were sufficient to learn to compensate for the fields themselves. This is unexpected since humans and monkeys can use color cues to perform a wide range of visuomotor tasks. Here we show that a monkey with extensive practice compensating for viscous fields in an elbow-movement task can use associated color cues to adjust motor output in anticipation of an impending field before physically encountering it.

Adaptation, Physiological↗

Simultaneous encoding of multiple potential reach directions in dorsal premotor cortex.

We present evidence that the primate brain can simultaneously generate discrete directional signals related to multiple alternative reaching actions before making a decision between them. A monkey performed a task in which the correct target for a reaching movement was specified during two consecutive instructed-delay periods. First, two potential targets were presented; and second, a nonspatial cue identified one of them as the correct movement target. During the first period, two directional signals coexisted in the activity of cells in dorsal premotor cortex (PMd), oriented toward the two potential targets. During the second period, one of these disappeared and the remaining signal predicted the monkey's response choice. These results suggest that, when faced with multiple salient opportunities for reaching, the primate brain performs sensorimotor transformations in parallel to begin planning several reaching movements simultaneously before selecting one for overt execution.

Animals↗

Modest gaze-related discharge modulation in monkey dorsal premotor cortex during a reaching task performed with free fixation.

Recent studies have shown that gaze angle modulates reach-related neural activity in many cortical areas, including the dorsal premotor cortex (PMd), when gaze direction is experimentally controlled by lengthy periods of imposed fixation. We looked for gaze-related modulation in PMd during the brief fixations that occur when a monkey is allowed to look around freely without experimentally imposed gaze control while performing a center-out delayed arm-reaching task. During the course of the instructed-delay period, we found significant effects of gaze angle in 27-51% of PMd cells. However, for 90-95% of cells, these effects accounted for <20% of the observed discharge variance. The effect of gaze was significantly weaker than the effect of reach-related variables. In particular, cell activity during the delay period was more strongly related to the intended movement expressed in arm-related coordinates than in gaze-related coordinates. Under the same experimental conditions, many cells in medial parietal cortex exhibited much stronger gaze-related modulation and expressed intended movement in gaze-related coordinates. In summary, gaze direction-related modulation of cell activity is indeed expressed in PMd during the brief fixations that occur in natural oculomotor behavior, but its overall effect on cell activity is modest.

Action Potentials↗