We investigated the idea that the cerebellum is required for precise timing of fast skilled arm movements by studying one situation where timing precision is required, namely finger opening in overarm throwing. Specifically, we tested the hypothesis that in overarm throws made by cerebellar patients, ball high-low inaccuracy is due to disordered timing of finger opening. Six cerebellar patients and six matched control subjects were instructed to throw tennis balls at three different speeds from a seated position while angular positions in three dimensions of five arm segments were recorded at 1,000 Hz with the search-coil technique. Cerebellar patients threw more slowly than controls, were markedly less accurate, had more variable hand trajectories, and showed increased variability in the timing, amplitude, and velocity of finger opening. Ball high-low inaccuracy was not related to variability in the height or direction of the hand trajectory or to variability in finger amplitude or velocity. Instead, the cause was variable timing of finger opening and thereby ball release occurring on a flattened arc hand trajectory. The ranges of finger opening times and ball release times (timing windows) for 95% of the throws were on average four to five times longer for cerebellar patients; e.g., across subjects mean ball release timing windows for throws made under the medium-speed instruction were 11 ms for controls and 55 ms for cerebellar patients. This increased timing variability could not be explained by disorder in control of force at the fingers. Because finger opening in throwing is likely controlled by a central command, the results implicate the cerebellum in timing the central command that initiates finger opening in this fast skilled multijoint arm movement.
1. While making saccades between targets with the head stationary, eye positions are constrained to two of the possible three degrees of freedom. Classically this constraint has been described by Donders' and Listing's laws. The objective was to determine whether these laws also apply for the straight arm when pointing between different targets. Thus we determined whether the arm adopts only one angular position for every pointing direction (Donders' law) and whether these positions can be described by rotations from a reference position about axes that lie in a plane (Listing's law). 2. The angular positions (orientations) of the arm in three-dimensional space were studied as subjects pointed with a straight arm at different targets. Arm position was measured with the search coil technique by means of coils attached to the back of the hand. Pointing was studied over a range of +/- 45 degrees in all directions from a central target located 45 degrees to the right of the straight-ahead position. 3. The positions of the arm in space were described by quaternion vectors, i.e., a particular position was described in terms of the axis and amplitude of a rotation from a reference position to that position. Using this description, it was found that the straight arm adopted a similar orientation (standard deviations ranged from 2.8 to 4.8 degrees) when pointing at a particular target irrespective of which target from which it had moved. 4. The angular position vectors for arm positions associated with relatively small movements (e.g., less than +/- 30 degrees) lay in a flat surface with minimal torsion. At first sight, this surface appeared to be similar to Listing's plane of the eye. However, for positions associated with larger movements (e.g., +/- 45 degrees) it became apparent that, unlike the eye, the surface deviated from one obeying Listing's law, i.e., it was twisted and showed torsion like that produced by rotations around the horizontal and vertical axes of a Fick gimbal. (The characteristic of a Fick gimbal is that the vertical axis is fixed, whereas the horizontal axis moves with the gimbal.) 5. Although there were differences between subjects, all showed a twisted position vector surface. The twist was always in the same direction, and it was always less than that of a Fick gimbal. 6. This position vector surface had a similar shape whether the arm was stationary or was moving between targets, whether subjects pointed with or without vision, and whether the pointing arm had moved between targets or from a bent-elbow position on the lap.(ABSTRACT TRUNCATED AT 400 WORDS)
1. Accurate overarm throwing requires precise control of joint rotations so that the ball is released at the appropriate time on the appropriate hand trajectory. Inaccuracy in throws, in turn, must result from errors in the control of joint rotations. But do high and low throws result from disorders in the joint rotations that produce the hand trajectory or in those that cause ball release? Are they due to error at a particular joint or to accumulation of errors across a number of joints? The objective was to answer these questions and thereby to gain insight into the CNS control of joint rotations in a skilled arm movement task. 2. Ten subjects--male, right-handed recreational ball players, all accurate throwers--sat with a fixed trunk and threw tennis balls at a 9 x 9 grid of 6-cm target squares 1.5 or 3 m away. Rotations of five arm segments in three dimensions were measured at 1,000 Hz with the magnetic-field search-coil technique. Hand trajectory (translation) was computed from these rotations. 3. The cause of ball high-low inaccuracy was investigated by determining its relation with hand kinematic parameters that could potentially affect it. No statistically significant relation was found between height of ball impact on the target and height of the hand trajectory. In contrast, statistically significant relations appeared between height of ball impact on the target and both hand trajectory length at ball release (for 8 of 10 subjects) and finger and hand orientation in space at ball release (for all 10 subjects). 4. Three hypotheses were proposed to explain the variable finger and hand orientations in space at ball release, i.e., that they resulted from errors in velocity of rotation at one or more proximal joints (wrist, elbow, shoulder), timing of onset of rotation at one or more proximal joints, or timing of ball release (due to incorrect velocity or timing of onset of finger opening). All three mechanisms could result in inappropriate finger and hand orientations in space at ball release, but the pattern of joint space trajectories would be different in each case. 5. High and low throws did not follow the joint space paths predicted by the first two hypotheses. Instead, as predicted by the third hypothesis, a separation of traces occurred when finger extension was plotted against wrist flexion or against elbow extension, e.g., for a given amplitude of wrist flexion, finger extension was large for the high throws and small for the low throws. 6. In agreement, when all throws were considered, a statistically significant (P < 0.005) relation was found between ball impact height on the target and the amplitude of finger extension, for a fixed amplitude of wrist flexion (10 subjects), and for a fixed amplitude of elbow extension (8 subjects). Only two subjects showed a statistically significant relation between ball impact height and the amplitude of wrist flexion, for a fixed amplitude of elbow extension. 7. The separation of finger extension-wrist flexion traces in joint space for high and low throws was ...
Previous studies have indicated that timing of finger opening in an overarm throw is likely controlled centrally, possibly by means of an internal model of hand trajectory. The present objective was to extend the study of throwing to an examination of the dynamics of finger opening. Throwing a heavy ball and throwing a light ball presumably require different neural commands, because the weight of the ball affects the mechanics of the arm, and particularly, the mechanics of the finger. Yet finger control is critical to the accuracy of an overarm throw. We hypothesized that finger opening in an overarm throw is controlled by a central mechanism that uses an internal model to predict and compensate for movement-dependent back forces on the fingers. To test this idea we determined whether finger motion is affected by back forces, i.e., whether larger back forces cause larger finger extensions. Back forces were varied by having subjects throw, at the same fast speed, tennis-sized balls of different weights (14, 55, and 196 g). Arm- and finger-joint rotations were recorded with the search-coil technique; forces on the middle finger were measured with force transducers. Recordings showed that during ball release, the middle finger experienced larger back forces in throws with heavier balls. Nevertheless, most subjects showed proximal interphalangeal joint extensions that were unchanged or actually smaller with the heavier balls. This was the case for the first throw and for all subsequent throws with a ball of a new weight. This suggests that the finger flexors compensated for the larger back forces by exerting larger torques during finger extension. Supporting this view, at the moment of ball release, all finger joints flexed abruptly due to the now unopposed torques of the finger flexors, and the amplitude of this flexion was proportional to ball weight. We conclude that in overarm throws made with balls of different weights, the CNS predicts the different back forces from the balls and adjusts finger flexor torques accordingly. This is consistent with the view that finger opening in overarm throws is controlled by means of an internal model of the motor apparatus and the external load.
1. Overarm throws made with the nondominant arm are usually less accurate than those made with the dominant arm. The objective was to determine the errors in the joint rotations associated with this inaccuracy, and thereby to gain insight into the neural mechanisms that contribute to skill in overarm throwing. 2. Overarm throws from both left and right arms were recorded on different occasions as six right-handed subjects sat with a fixed trunk and threw 150 tennis balls at about the same speed at a 6-cm square on a target grid 3 m away. Joint rotations at the shoulder, elbow, wrist, and finger, and arm translations, were computed from recordings of arm segment orientations made with the magnetic-field search-coil technique. 3. All subjects threw less accurately in this task with the left (nondominant) arm. For throws made with the left arm, the height of ball impact on the target grid was related to hand trajectory length and to hand orientation in space at ball release, but not to hand trajectory height. 4. Two hypotheses were proposed to explain the decreased ball accuracy in the high-low direction during throwing with the nondominant arm: that it was caused by increased variability in the velocity or timing of onset of rotations at proximal joints (which determine the path of the hand through space) or increased variability in the velocity or timing of onset of finger extension (which determine the moment of ball release). 5. A prediction of the first hypothesis was that proximal joint rotations should be more variable in throws with the left arm. This was the case for the majority of proximal joint rotations in the six subjects when variability was examined in joint space. However, some proximal joint rotations were more variable in the right arm. 6. The first hypothesis was directly tested by determining whether hand angular position in space (which represents the sum of all proximal joint rotations) was related to ball impact height on the target grid at a fixed translational position in the throw. No relation was found between these variables for throws with the left arm in four subjects, whereas a weak relation was found for two subjects. It was concluded that, considering all subjects, the first hypothesis could not explain the results. 7. In contrast, in agreement with the second hypothesis, a strong relation (P < 0.001) was found in all subjects between ball impact height on the target grid and time of ball release for throws with the left arm, and with time of onset of finger extension. 8. Across all six subjects the timing precision (windows) for 95% of the throws was (for ball release) right arm, 9.3 ms; left arm, 22.5 ms; (for onset of finger extension) right arm, 13.7 ms; left arm, 26.7 ms. 9. Timing of onset of finger extension was no less accurate than timing of onset of other joint rotations for both left and right arms. However, simulations of throws showed that, for the same error in timing, finger extension had twice as large an effect on ball direction as any other joint rotation. Timing e...
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