Miya K. Rand

Parallel neural networks for learning sequential procedures

Recent studies have shown that multiple brain areas contribute to different stages and aspects of procedural learning. On the basis of a series of studies using a sequence-learning task with trial-and-error, we propose a hypothetical scheme in which a sequential procedure is acquired independently by two cortical systems, one using spatial coordinates and the other using motor coordinates. They are active preferentially in the early and late stages of learning, respectively. Both of the two systems are supported by loop circuits formed with the basal ganglia and the cerebellum, the former for reward-based evaluation and the latter for processing of timing. The proposed neural architecture would operate in a flexible manner to acquire and execute multiple sequential procedures.

Differential roles of monkey striatum in learning of sequential hand movement.

Abstract To study the role of the basal ganglia in learning of sequential movements, we trained two monkeys to perform a sequential button-press task (2×5 task). This task enabled us to examine the process of learning new sequences as well as the execution of well-learned sequences repeatedly. We injected muscimol (a GABA agonist) into different parts of the striatum to inactivate the local neural activity reversibly. The learning of new sequences became deficient after injections in the anterior caudate and putamen, but not the middle-posterior putamen. The execution of well-learned sequences was disrupted after injections in the middle-posterior putamen and, less severely, after injections in the anterior caudate/putamen. These results suggest that the anterior and posterior portions of the striatum participate in different aspects of learning of sequential movements.

Movement accuracy constraints in Parkinson's disease patients.

This study examined the hypothesis that the kinematics of movements performed by PD (Parkinsons disease) patients are differentially affected depending on whether or not the aiming movement has an accuracy constraint. The aiming movements required elbow extension in the horizontal plane on a digitizer. There were two movement conditions: (1) one having a spatial accuracy requirement in which the subjects moved to the defined target and stopped on it; and (2) one requiring the subjects to move toward the defined target without stopping precisely on it. Subjects were instructed to make their movements as fast and as accurate as possible in response to the auditory imperative signal. PD patients modified the movement speed and kinematics depending on the two accuracy conditions. However, when the accuracy constraint was imposed, movement slowness observed in the patients was much more pronounced. The most revealing result was localized to the deceleration phase, particularly as the target was approached. The patients also were found to make a higher number of acceleration zero crossings from negative to positive to reach the target, indicating that the movements were more irregular. For the patients, the first acceleration zero crossing from negative to positive occurred much earlier in the movement than that for the controls. In addition, when movement accuracy was constrained, the number of zero crossings was accentuated. These data show that when PD patients make aiming movements to a target, their deceleration phase becomes longer and more variable.

Characteristics of a long-term procedural skill in the monkey

Abstract The purpose of this study was to characterize the nature and structure of procedural memory. We have previously studied the process of learning sequential behavioral procedures using monkeys. The monkey’s task was to press five consecutive pairs of buttons (indicated by illumination) in the correct order for every pair, which he had to find by trial-and-error in a block of trials. The whole sequence was called a “hyperset”; each pair was called a “set”. We first examined whether monkeys learned to perform a hyperset as a single sequence or learned the order of button-presses individually for each set. To answer this question, we generated hypersets that were the same as the hypersets that had been extensively learned except that the order of the sets was reversed. The performance of these “reversed hypersets” was much worse than the performance of the original learned hypersets and was similar to the performance of new hypersets, as regards both the number of errors and the performance time. The result suggests that monkeys learned a hyperset as a sequence. To examine whether the learned performance was specific to the hand used for practice, we had monkeys use the same hand throughout the long-term practice of each hyperset, and then tested the opposite hand. The performance using the opposite hand was worse than the performance using the trained hand, but was better than the performance for new hypersets. This indicates that the memory for the sequential procedure is only partially accessible to the hand that was not used for the practice.

EMG analysis of lower limb muscles in humans during quick change in running directions

Open and cross maneuvers for changing running direction were studied to characterize selective EMG activity between the maneuvers. Eleven subjects turned towards the right or the left during running. The gluteus medius modified foot trajectory of the leading leg during the open maneuver, whereas the sartorius worked modestly during the cross maneuver. Compared with the cross maneuver the open maneuver exhibited greater vastus medialis and gastrocnemius activity during the ground support phase, faster running speed and wider turning angle. These results suggest that the open maneuver is more effective than the cross maneuver for quickly changing running direction.

Effects of accuracy constraints on reach-to-grasp movements in cerebellar patients.

Reach-to-grasp movements of patients with pathology restricted to the cerebellum were compared with those of normal controls. Two types of paradigms with different accuracy constraints were used to examine whether cerebellar impairment disrupts the stereotypic relationship between arm transport and grip aperture and whether the variability of this relationship is altered when greater accuracy is required. The movements were made to either a vertical dowel or to a cross bar of a small cross. All subjects were asked to reach for either target at a fast but comfortable speed, grasp the object between the index finger and thumb, and lift it a short distance off the table. In terms of the relationship between arm transport and grip aperture, the control subjects showed a high consistency in grip aperture and wrist velocity profiles from trial to trial for movements to both the dowel and the cross. The relationship between the maximum velocity of the wrist and the time at which grip aperture was maximal during the reach was highly consistent throughout the experiment. In contrast, the time of maximum grip aperture and maximum wrist velocity of the cerebellar patients was quite variable from trial to trial, and the relationship of these measurements also varied considerably. These abnormalities were present regardless of the accuracy requirement. In addition, the cerebellar patients required a significantly longer time to grasp and lift the objects than the control subjects. Furthermore, the patients exhibited a greater grip aperture during reach than the controls. These data indicate that the cerebellum contributes substantially to the coordination of movements required to perform reach-to-grasp movements. Specifically, the cerebellum is critical for executing this behavior with a consistent, well-timed relationship between the transport and grasp components. This contribution is apparent even when accuracy demands are minimal.

Role of vision in aperture closure control during reach-to-grasp movements

We have previously shown that the distance from the hand to the target at which finger closure is initiated during the reach (aperture closure distance) depends on the amplitude of peak aperture, as well as hand velocity and acceleration. This dependence suggests the existence of a control law according to which a decision to initiate finger closure during the reach is made when the hand distance to target crosses a threshold that is a function of the above movement-related parameters. The present study examined whether the control law is affected by manipulating the visibility of the hand and the target. Young adults made reach-to-grasp movements to a dowel under conditions in which the target or the hand or both were either visible or not visible. Reaching for and grasping a target when the hand and/or target were not visible significantly increased transport time and widened peak aperture. Aperture closure distance was significantly lengthened and wrist peak velocity was decreased only when the target was not visible. Further analysis showed that the control law was significantly different between the visibility-related conditions. When either the hand or target was not visible, the aperture closure distance systematically increased compared to its value for the same amplitude of peak aperture, hand velocity, and acceleration under full visibility. This implies an increase in the distance-related safety margin for grasping when the hand or target is not visible. It has been also found that the same control law can be applied to all conditions, if variables describing hand and target visibility were included in the control law model, as the parameters of the task-related environmental context, in addition to the above movement-related parameters. This suggests that that the CNS utilizes those variables for controlling grasp initiation based on a general control law.

Adaptive changes in responses to repeated locomotor perturbations in cerebellar patients

Abstract This study examined the responses of cerebellar patients and a group of age- and sex-matched control subjects to repeated changes in treadmill speed in order to test whether cerebellar patients can adapt their gait to this type of perturbation and, if so, whether their responses are comparable to those of controls. While the subject walked on the treadmill, a perturbation consisting of a sudden slowing of the treadmill followed by a sudden increase back to the original speed was applied repeatedly at a specific time during the step cycle. Both the control subjects and cerebellar patients were able to compensate for the perturbations by minimizing their postural sway and changing step length. However, the nature of the compensatory changes in step length differed between these subject groups. Control subjects compensated for the perturbation by consistently using the same leg to initiate the response to the perturbation and by adapting a pattern of stepping such that the EMG characterizing the response occurred in a manner that was entrained to the timing of the normal locomotor cycle. In contrast, the patients, although undergoing modifications in step length, employed a much less consistent motor pattern from trial to trial than that of the normal subjects. An inconsistent pattern among their responses was apparent in both the analysis of stepping and in the EMG activity of the gastrocnemius and anterior tibial muscles. These results suggest that, although the cerebellar patients can adapt their behavior in response to locomotor perturbations, they do not establish a motor pattern comparable to that employed by normal subjects.

The influence of movement segment difficulty on movements with two-stroke sequence

Abstract Arm movements in the horizontal plane consisting of two segments were examined to determine whether the difficulty of the second segment influenced the kinematic characteristics of the first segment. The direction of the first segment was an elbow extension movement away from the trunk and remained constant throughout the experiment. The direction of the second segment varied between forearm extension and flexion movements. Based on Fitts’ law, two different indexes of difficulty (ID) of the second segment were utilized by changing target size and movement amplitude. The effects of changing ID were examined for two different movement amplitudes. All movements were single-joint movements employing elbow flexion/extension and were recorded by an x-y digitizer. Variations in the ID of the second segment produced context-dependent kinematic changes in the performance of the initial segment. Movement duration increased when the ID was increased by reducing target size for both extension-extension sequence and extension-flexion sequences. Peak velocity also decreased for higher ID targets in the extension-flexion sequence. However, there was an interaction between the ID and movement amplitude in the extension-flexion sequence. In this sequence the duration of movement for the high ID/large movement amplitude condition increased substantially compared with the low ID/small movement amplitude condition. In addition, changing ID of the second segment influenced the time between the two segments (intersegment interval) in the extension-flexion sequence. Collectively, these data suggest that the planning of complex movements is based in part on the accuracy demands of multiple segments of the sequence.

Control of aperture closure during reach-to-grasp movements in parkinson’s disease

This study examined whether the pattern of coordination between arm-reaching toward an object (hand transport) and the initiation of aperture closure for grasping is different between PD patients and healthy individuals, and whether that pattern is affected by the necessity to quickly adjust the reach-to-grasp movement in response to an unexpected shift of target location. Subjects reached for and grasped a vertical dowel, the location of which was indicated by illuminating one of the three dowels placed on a horizontal plane. In control conditions, target location was fixed during the trial. In perturbation conditions, target location was shifted instantaneously by switching the illumination to a different dowel during the reach. The hand distance from the target at which the subject initiated aperture closure (aperture closure distance) was similar for both the control and perturbation conditions within each group of subjects. However, that distance was significantly closer to the target in the PD group than in the control group. The timing of aperture closure initiation varied considerably across the trials in both groups of subjects. In contrast, aperture closure distance was relatively invariant, suggesting that aperture closure initiation was determined by spatial parameters of arm kinematics rather than temporal parameters. The linear regression analysis of aperture closure distance showed that the distance was highly predictable based on the following three parameters: the amplitude of maximum grip aperture, hand velocity, and hand acceleration. This result implies that a control law, the arguments of which include the above parameters, governs the initiation of aperture closure. Further analysis revealed that the control law was very similar between the subject groups under each condition as well as between the control and perturbation conditions for each group. Consequently, the shorter aperture closure distance observed in PD patients apparently is a result of the hypometria of their grip aperture and bradykinesia of hand transport movement, rather than a consequence of a deficit in transport-grasp coordination. It is also concluded that the perturbation of target location does not disrupt the transport-grasp coordination in either healthy individuals or PD patients.