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Biomedical subjects

C Ghez

Publications and source records attributed to C Ghez.

At least 19 recordsLinked to original sources

Kinematic analysis of reaching in the cat.

The present study examines the kinematic features of forelimb movements made by cats reaching for food in horizontal target wells located at different heights and distances. Wrist paths consisted of two relatively straight segments joined at a "via-point" in front of the aperture of the food well. In the initial lift phase, the paw was raised to the via-point in front of the target. In the second, or thrust phase, the paw was directed forward into the food well. During the lift, the paw was moved toward the target primarily by elbow flexion, accompanied by a sequence of biphasic shoulder and wrist movements. Thrust was accomplished primarily by shoulder flexion while the wrist and the paw were maintained at near-constant angles. The animals varied the height of the reach primarily by varying elbow flexion with proportional changes in elbow angular velocity and angular acceleration and with corresponding variations in wrist speed. Thus, cats reached for targets at different heights by scaling a common kinematic profile. Over a relatively large range of target heights, animals maintained movement duration constant, according to a simple "pulse-height" control strategy (isochronous scaling). For reaches to a given target height, animals compensated for variability in peak acceleration by variations in movement time. We examined the coordination between the shoulder and the wrist with the elbow. Early during the lift, peak shoulder extensor and peak elbow flexor accelerations were synchronized. Late during the lift phase, wrist extensor acceleration was found to occur during the period of elbow flexor deceleration. We hypothesize that these linkages could, in part, be due to passive mechanical interactions. To determine how the angular trajectories of the different joints were organized in relation to target location, we plotted joint kinematic changes directly on the wrist and MCP joint paths. These plots revealed that for all target heights and movement speeds, wrist extensor deceleration occurred at approximately the same spatial location with respect to the target. This analysis also demonstrated that the second phase of MCP flexion occurred when the paw was below the lower lip of the food well, while the subsequent extension occurred after the tip cleared this obstacle. During thrust, wrist and MCP angles were maintained, reflecting the need to align the paw within the food well. Our findings suggest that cats plan the reaching phase of prehension as a sequence of discrete movement segments, each serving a particular goal in the task, rather than as an single unit.

Animals

Proprioceptive control of interjoint coordination.

This paper reviews a series of experiments comparing intact controls with functionally deafferented patients to determine the role of proprioception in controlling dynamic interactions between limb segments during movement. We examine the control of hand path in a planar movement-reversal task and in a familiar three-dimensional gesture with similar biomechanical characteristics. In the planar task subjects had to move their hand out and back along a series of straight-line segments in the horizontal plane without visual feedback. The lengths and directions of the target line segments were chosen to require different amounts of shoulder motion while requiring the same elbow excursion. In controls, hand paths were, as required, straight with sharp bends at the outermost point. In patients, however, distinctive errors appeared at movement reversals, consisting of widened hand paths resulting from desynchronization in the reversals of elbow and shoulder motions. These errors reflected an inability to program elbow muscle contractions in accord with interaction torques produced at the elbow by variations in acceleration of the shoulder. The reversal errors were substantially reduced after patients had practiced for a few trials while visually monitoring movements of their arm. The improvement was not limited to the direction where they had practiced with vision, but also extended to other directions in which the elbow torques were different. This suggests that practice with vision of the arm served to improve the general rules that subjects used to plan movement, rather than simply improving the performance of a specific response. Similar to their performance on the planar task, the patients made errors in interjoint coordination during unconstrained three-dimensional gestures with movement reversals. We conclude (i) that both the planning and the learning of movement required an internal model of the dynamic properties of the limb that takes account of interaction torques acting at different joints; (ii) that this internal model is normally established and updated using proprioceptive information; but (iii) that when proprioception is lacking, vision of the limb in motion partially substitutes for proprioception.

Adult

Impairments of reaching movements in patients without proprioception. I. Spatial errors.

1. This paper introduces a series of studies in which we analyze the impairments in a planar reaching task in human patients with severe proprioceptive deficits resulting from large-fiber sensory neuropathy. We studied three patients, all of whom showed absence of discriminative tactile sensation, position sense, and stretch reflexes in the upper extremities. Muscle strength was normal. We compared the reaching movements of the patients with those of normal control subjects. The purpose of this first paper was no characterize the spatial errors in these patients that result primarily from impairments in the planning and execution of movement rather than in feedback control. This was done by using a task in which visual feedback of errors during movement was prevented. 2. Subjects were instructed to move their hand from given starting positions of different targets on a horizontal digitizing tablet. Hand position and targets were displayed on a computer screen. Subjects could not see their hand, and the screen display of hand position was blanked at the signal to move. Thus visual feedback during movement could not be used to achieve accuracy. Movement paths were displayed as knowledge of results after each trial. 3. Compared with controls, the patients made large spatial errors in both movement direction and extent. Directional errors were evident from movement onset, suggesting that they resulted from improper planning. In addition, patients' hand paths showed large curves and secondary movements after initial stops. 4. The overall control strategy used by patients appeared the same as that used by controls. Hand trajectories were approximately bell shaped, and movement extent was controlled by scaling a trajectory waveform in amplitude and time. However, both control subjects and patients showed systematic errors in movement extent that depended on the direction of hand movement. In control subjects, these systematic dependencies of extent on direction were small, but in patients they produced large and prominent errors. Analysis of the hand trajectories revealed that errors were associated with differences in velocity and acceleration for movements in different directions. In an earlier study, we showed that in subjects with normal sensation that the dependence of acceleration and velocity on direction results from a failure to take the inertial properties of the limb into account in programming the initial trajectory. In control subjects, these differences in initial acceleration are partially compensated by direction-dependent variations in movement time.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult

Impairments of reaching movements in patients without proprioception. II. Effects of visual information on accuracy.

1. The aim of this study was to determine how vision of a cursor indicating hand position on a computer screen or vision of the limb itself improves the accuracy of reaching movements in patients deprived of limb proprioception due to large-fiber sensory neuropathy. In particular, we wished to ascertain the contribution of such information to improved planning rather than to feedback corrections. We analyzed spatial errors and hand trajectories of reaching movements made by subjects moving a hand-held cursor on a digitizing tablet while viewing targets displayed on a computer screen. The errors made when movements were performed without vision of their arm or of a screen cursor were compared with errors made when this information was available concurrently or prior to movement. 2. Both monitoring the screen cursor and seeing their limb in peripheral vision during movement improved the accuracy of the patients' movements. Improvements produced by seeing the cursor during movement are attributable simply to feedback corrections. However, because the target was not present in the actual workspace, improvements associated with vision of the limb must involve more complex corrective mechanisms. 3. Significant improvements in performance also occurred in trials without vision that were performed after viewing the limb at rest or during movements. In particular, prior vision of the limb in motion improved the ability of patients to vary the duration of movements in different directions so as to compensate for the inertial anisotropy of the limb. In addition, there were significant reductions in directional errors, path curvature, and late secondary movements. Comparable improvements in extent, direction, and curvature were produced when subjects could see the screen cursor during alternate movements to targets in different directions. 4. The effects of viewing the limb were transient and decayed during a period of minutes once vision of the limb was no longer available. 5. It is proposed that the improvements in performance produced after vision of the limb were mediated by the visual updating of internal models of the limb. Vision of the limb at rest may provide configuration information while vision of the limb in motion provides additional dynamic information. Vision of the cursor and the resulting ability to correct ongoing movements, however, is considered primarily to provide information about the dynamic properties of the limb and its response to neural commands.

Adult

Control of limb dynamics in normal subjects and patients without proprioception.

1. We recently showed that patients lacking proprioceptive input from their limbs have particular difficulty performing multijoint movements. In a pantomimed slicing gesture requiring sharp reversals in hand path direction, patients showed large hand path distortions at movement reversals because of failure to coordinate the timing of the separate reversals at the shoulder and elbow joints. We hypothesized that these reversal errors resulted from uncompensated effects of inertial interactions produced by changes in shoulder joint acceleration that were transferred to the elbow. We now test this hypothesis and examine the role of proprioceptive input by comparing the motor performance of five normal subjects with that of two patients with large-fiber sensory neuropathy. 2. Subjects were to trace each of six template lines presented randomly on a computer screen by straight overlapping out-and-back movements of the hand on a digitizing tablet. The lines originated from a common starting position but were in different directions and had different lengths. Directions and lengths were adjusted so that tracing movements would all require the same elbow excursion, whereas shoulder excursion would vary. The effects of varying interaction torques on elbow kinematics were then studied. The subject's dominant arm was supported in the horizontal plane by a low-inertia brace equipped with ball bearing joints and potentiometers under the elbow and shoulder. Hand position was monitored by a magnetic pen attached to the brace 1 cm above a digitizing tablet and could be displayed as a screen cursor. Vision of the subject's arm was blocked and the screen cursor was blanked at movement onset to prevent visual feedback during movement. Elbow joint torques were calculated from joint angle recordings and compared with electromyographic recordings of elbow joint musculature. 3. In control subjects, outward and inward paths were straight and overlapped the template lines regardless of their direction. As prescribed by the task, elbow kinematics remained the same across movement directions, whereas interaction torques varied substantially. The timing of the onsets of biceps activity and the offsets of triceps activity during elbow flexion varied systematically with direction-dependent changes in interaction torques. Controls exploited or dampened these interaction torques as needed to meet the kinematic demands of the task. 4. In contrast, the patients made characteristic errors at movement reversals that increased systematically across movement directions. These reversal errors resulted from improper timing of elbow and shoulder joint reversals.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult

Learning a visuomotor transformation in a local area of work space produces directional biases in other areas.

1. The dependence of directional biases in reaching movements on the initial position of the hand was studied in normal human subjects moving their unseen hand on a horizontal digitizing tablet to visual targets displayed on a vertical computer screen. 2. When initial hand positions were to the right of midline, movements were systematically biased clockwise. Biases were counterclockwise for starting points to the left. Biases were unaffected by the screen location of the starting and target positions. 3. Vision of the hand in relation to the target before movement, as well as practice with vision of the cursor during the movement, temporarily eliminated these biases. The spatial organization of the biases suggests that, without vision of the limb, the nervous system underestimates the distance of the hand from an axis or plane that includes its most common operating location. 4. To test the hypothesis that such an underestimate might represent an adaptation to a local area of work space or range effect, subjects were trained to reach accurately from right or left positions. After training, movements initiated from other locations, including ones that were previously error free, showed new biases that again represented underestimates of the distance of the initial hand position from the new trained location. 5. We conclude that hand path planning is dependent on learned representations of the location of the hand in the work space.

Adult

Coherent modulations of human motor unit discharges during quasi-sinusoidal isometric muscle contractions.

Spectral analysis of single-unit discharges, multi-unit EMG and muscle force during voluntary quasi-sinusoidal isometric contractions of two hand muscles revealed corresponding modulations of the firing rates of motor units at the frequency of the force oscillation. These rate modulations were correlated; and they showed a phase advance over the force oscillation, which is consistent with a cause-effect relationship between changes in firing rate and variations in force. These effects, observed over wide ranges of modulation amplitudes and frequencies, confirm the role of rate coding in the generation of time-varying muscle contractions; and they support the idea that during voluntary contraction of a given muscle, the motoneuron pool is subject to a common drive.

Adult

Accuracy of planar reaching movements. I. Independence of direction and extent variability.

This study examined the variability in movement end points in a task in which human subjects reached to targets in different locations on a horizontal surface. The primary purpose was to determine whether patterns in the variable errors would reveal the nature and origin of the coordinate system in which the movements were planned. Six subjects moved a hand-held cursor on a digitizing tablet. Target and cursor positions were displayed on a computer screen, and vision of the hand and arm was blocked. The screen cursor was blanked during movement to prevent visual corrections. The paths of the movements were straight and thus directions were largely specified at the onset of movement. The velocity profiles were bell-shaped, and peak velocities and accelerations were scaled to target distance, implying that movement extent was also programmed in advance of the movement. The spatial distributions of movement end points were elliptical in shape. The major axes of these ellipses were systematically oriented in the direction of hand movement with respect to its initial position. This was true for both fast and slow movements, as well as for pointing movements involving rotations of the wrist joint. Using principal components analysis to compute the axes of these ellipses, we found that the eccentricity of the elliptical dispersions was uniformly greater for small than for large movements: variability along the axis of movement, representing extent variability, increased markedly but nonlinearly with distance. Variability perpendicular to the direction of movement, which results from directional errors, was generally smaller than extent variability, but it increased in proportion to the extent of the movement. Therefore, directional variability, in angular terms, was constant and independent of distance. Because the patterns of variability were similar for both slow and fast movements, as well as for movements involving different joints, we conclude that they result largely from errors in the planning process. We also argue that they cannot be simply explained as consequences of the inertial properties of the limb. Rather they provide evidence for an organizing mechanism that moves the limb along a straight path. We further conclude that reaching movements are planned in a hand-centered coordinate system, with direction and extent of hand movement as the planned parameters. Since the factors which influence directional variability are independent of those that influence extent errors, we propose that these two variables can be separately specified by the brain.

Adult

Accuracy of planar reaching movements. II. Systematic extent errors resulting from inertial anisotropy.

This study examines the source of direction-dependent errors in movement extent made by human subjects in a reaching task. As in the preceding study, subjects were to move a cursor on a digitizing tablet to targets displayed on a computer monitor. Movements were made without concurrent visual feedback of cursor position, but movement paths were displayed on the monitor after the completion of each movement. We first examined horizontal hand movements made at waist level with the upper arm in a vertical orientation. Targets were located at five distances and two directions (30 degrees and 150 degrees) from one of two initial positions. Trajectory shapes were stereotyped, and movements to more distant targets had larger accelerations and velocities. Comparison of movements in the two directions showed that in the 30 degrees direction responses were hypermetric, accelerations and velocities were larger, and movement times were shorter. Since movements in the 30 degrees direction required less motion of the upper arm than movements in the 150 degrees direction, we hypothesized that the differences in accuracy and acceleration reflected a failure to take into account the difference in total limb inertia in the two directions. To test this hypothesis we simulated the initial accelerations of a two-segment limb moving in the horizontal plane with the hand at shoulder level when a constant force was applied at the hand in each of 24 directions. We compared these simulated accelerations to ones produced by our subjects with their arms in the same position when they aimed movements to targets in the 24 directions and at equal distances from an initial position. The magnitudes of both simulated and actual accelerations were greatest in the two directions perpendicular to the forearm, where inertial resistance is least, and lowest for movements directed along the axis of the forearm. In all subjects, the directional variation in peak acceleration was similar to that predicted by the model and shifted in the same way when the initial position of the hand was displaced. The pattern of direction-dependent variations in initial acceleration did not depend on the speed of movement. It was also unchanged when subjects aimed their movements toward targets presented within the workspace on the tablet instead of on the computer monitor. These findings indicate that, in programming the magnitude of the initial force that will accelerate the hand, subjects do not fully compensate for direction dependent differences in inertial resistance.(ABSTRACT TRUNCATED AT 400 WORDS)

Acceleration

Differential effects of local inactivation within motor cortex and red nucleus on performance of an elbow task in the cat.

This study examined changes in the performance of a single-joint, elbow task produced by reversible inactivation of local regions within the proximal forelimb representation in area 4 gamma of motor cortex (MCx) and the red nucleus (RN) of the cat. Inactivation was carried out by microinjecting lidocaine, gamma-aminobutyric acid, or muscimol into sites where microstimulation evoked contraction of elbow muscles. Reaction time, amplitude, and speed (velocity or dF/dt) of position and force responses elicited during inactivation were compared to control values obtained immediately prior to inactivation. In addition, we assessed qualitatively the effects of inactivation on reaching, placing reactions, and proprioceptive responses to imposed limb displacement. In the single-joint task, injections in MCx did not increase reaction time (simple or choice) and produced modest and inconsistent reductions in response amplitude (mean -8%) and speed (mean -19%). In contrast, injections of the same amounts of inactivating agents in the forelimb representation of RN consistently increased reaction time (34.4%), and increased the reaction time coefficient of variability (32%). There were small reductions in response amplitude (-4%) and speed (-10%) which were less than those produced by MCx inactivation. During reaching, however, these same injections in MCx and RN produced a substantial loss of accuracy. For MCx, this was due, in part, to systematic hypometria: for RN, inaccuracy resulted from increased variability in paw paths. Placing reactions and corrective responses to imposed limb displacements were also depressed by the cortical and rubral injections. Our results suggest that the forelimb representation in RN plays a role in the initiation of the single-joint, elbow tracking response examined here. The RN may mediate cerebellar regulation of response timing, a function that is likely to be important for interjoint coordination. Although neurons in the forelimb representations of MCx may contribute to force generation in single-joint movements, their contribution to multijoint control appears to be more important and is examined in the subsequent report (Martin and Ghez 1993).

Animals

Differential impairments in reaching and grasping produced by local inactivation within the forelimb representation of the motor cortex in the cat.

This study analyzed changes in the performance of a reaching task and its adaptive modification produced by reversible inactivation of three sites within the forelimb representation of the motor cortex (MCx, area 4 gamma) in five cats by microinjections of muscimol. Two sites were located in the lateral MCx, rostral (RL-MCx) and caudal (CL-MCx) to the end of the cruciate sulcus, where intracortical microstimulation (ICMS) produced contraction of the most distal muscles. The third site was located more medially, in the anterior sigmoid gyrus (RM-MCx) where ICMS primarily produced contraction of more proximal muscles. The task required the animals to reach into a horizontal target well, located in front of them at one of three possible heights, to grasp and retrieve a small piece of food. The height of the reach was primarily achieved by elbow flexion. Grasping consisted primarily of digit flexion, and food retrieval consisted of forearm supination and shoulder extension. In some blocks of trials, an obstacle was placed in the path of the limb to assess the animal's ability to adaptively adjust the kinematic characteristics of their response trajectory. In normal animals, contact with the bar on the first trial triggered a corrective response at short latency that allowed the paw to circumvent the bar. On all subsequent trials, the trajectory was adapted to prevent contact with the obstacle, with a safety margin of about 1 cm. Inactivation at all sites produced a slowing of movement, a protracted and extended forelimb posture, and increased variability of initial limb position. In addition, inactivation of RL-MCx immediately produced systematic reaching errors, consisting of hypermetric movements, as well as impaired grasping and food retrieval. The degree of hypermetria was similar for all target heights and was not associated with alterations in trajectory control. During inactivation, animals did not compensate for the hypermetria by reducing paw path elevation, suggesting a defect in kinematic planning or in adaptive control. This was confirmed by finding that trajectory adaptation to avoid bar contact was impaired during RL-MCx inactivation. The short latency corrective response, triggered by contact of the limb with the obstacle was, however, preserved. Inactivation of CL-MCx did not impair aiming, grasping, or adaptation immediately after injection. However, impairments occurred after about 1 h postinjection, and at that time mimicked the effects of RL-MCx inactivation.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Loss of proprioception produces deficits in interjoint coordination.

1. We analyzed the performance of a simple pantomimed gesture in 2 patients with large-fiber sensory neuropathy and 11 control subjects to determine how proprioceptive deafferentation disrupts unconstrained multijoint movements. Both patients had near-total loss of joint position, vibration, and discriminative touch sensation in the upper extremities. Muscle strength remained intact. 2. Subjects performed a gesture similar to slicing a loaf of bread. In this gesture, the hand first moves outward from the body, reverses direction sharply, and then moves back toward the body. Accurate performance requires precise coordination between the shoulder and elbow joints during movement reversals. Movements were performed under two conditions: with eyes open and with eyes closed. Three dimensional shoulder, elbow, wrist, and hand trajectories were recorded on a WATSMART system. 3. When control subjects performed the gesture with their eyes closed, their wrist trajectories were relatively straight and individual cycles of motion were planar. Movements reversed direction sharply, such that outward and inward portions of the wrist path were closely aligned. Corresponding to this spatial profile, the reversals in movement direction at the shoulder joint, from flexion to extension, and at the elbow joint, from extension to flexion, were synchronous. 4. In contrast, when deafferented patients performed the gesture with their eyes closed, their wrist trajectories were highly curved and individual cycles were severely nonplanar. The wrist paths showed a characteristic anomaly during the reversal in movement direction, when elbow joint movement became transiently locked. Correspondingly, the movement reversals at the shoulder and elbow joints were severely temporally decoupled. 5. When patients were able to view their limbs during performance of this gesture there was significant improvement in the linearity and planarity of movements. However, the patients remained unable to synchronize the movements at the shoulder and elbow joints to produce spatially precise wrist paths. 6. We conclude that loss of proprioception disrupts interjoint coordination and discuss the hypothesis that this interjoint coordination deficit results from a failure to control the interaction forces that arise between limb segments during multijoint movements.

Adult

Impairments in reaching during reversible inactivation of the distal forelimb representation of the motor cortex in the cat.

We report changes in the performance of a prehension task in the cat following focal inactivation of the rostrolateral subregion of the distal forelimb area of motor cortex (MCx) produced by muscimol microinjection. Animals reached into a cylindrical target to retrieve a morsel of food. Movements consisted of distinct lift and forward thrust phases following which the food was grasped and retrieved. In separate blocks of trials an obstacle was inserted in the path of the limb. Impact evoked an immediate compensatory trajectory change to bypass the obstruction and, on subsequent trials, an adaptive trajectory change to avoid impact. Inactivation produced three major defects: (1) uncompensated aiming biases to a location above the target; (2) loss of coordination of the grasp and food retrieval; and (3) impairment in trajectory adaptation to avoid impact of the limb with an obstacle. Thus, focal inactivation of the distal forelimb area of MCx produced disordered control of all forelimb joints. The impairment in trajectory adaptation and failure to compensate for aiming biases suggests that the MCx is important in motor learning.

Animals

Task-related coding of stimulus and response in cat red nucleus.

In the present study we recorded the activity of single neurons in the forelimb area of red nucleus (RN) during performance of three step-tracking tasks designed to dissociate the coding of stimulus and response variables in the discharge of recorded neurons. In two of these tasks, the standard and stimulus-reversal arm tasks, elbow flexion and extension were elicited by different stimuli enabling us to distinguish activity correlated with the forelimb response from the stimulus eliciting it. The third task (neck task) allowed us to determine whether neuronal modulation was related to an unconditioned orienting response that occurred concurrently with the forelimb response. We have previously reported that these three tasks separate neurons in MCx whose modulation precedes the response (lead cells) into three distinct classes in which task-related activity either is correlated with the direction of the forelimb response, correlated with the stimulus, or not correlated with either (Martin and Ghez 1985). All lead cells, however, remained timed to the stimulus rather than to the response. The present results show that RN lead cells can be subdivided into the same three classes as those in MCx and their discharge was also contingent on the subsequent production of a behavioral response. (1) Force-direction neurons (35%; n = 16) showed changes in activity correlated with the production of forearm force in a particular direction suggesting that they could participate in selecting the appropriate forelimb response. The onset of task-related modulation of activity was better timed to the response, in contrast to force-direction neurons in MCx, which were better timed to the stimulus. (2) Stimulus-direction neurons (18%; n = 8) modulated their activity in relation to a particular stimulus evoking either flexor or extensor responses and during neck task performance. These neurons could be involved in processing stimulus information or in the production of neck torque. The task-related discharge of these lead cells was better timed to the stimulus than to either the forelimb or the neck response. (3) Nondirectional neurons (47%; n = 21) modulated their activity during all tasks examined. Their discharge did not correlate with any specific feature of the stimulus or response, and as a group, was better timed to the stimulus than to the response. Nondirectional neurons may participate in some aspect of motor preparation.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Organization of voluntary movement.

There have recently been a number of advances in our knowledge of the organization of complex, multi-joint movements. Promising starts have been made in our understanding of how the motor system translates information about the location of external targets into motor commands encoded in a body-based coordinate system. Two simplifying strategies for trajectory control that are discussed are parallel specification of response features and the programming of equilibrium trajectories. New insights have also been gained into how neural systems process sensory information to plan and assist with task performance. A number of recent papers emphasize the feedforward use of sensory input, which is mediated through models of the external world, the body's physical plant, and the task structure. These models exert their influence at both reflex and higher levels and permit the preparation of predictive default parameters of trajectories as well as strategies for resolving task demands.

Animals

Trajectory control in targeted force impulses. VII. Independent setting of amplitude and direction in response preparation.

We have previously shown that when aiming impulses of force to unpredictable flexion and extension targets, subjects prepare themselves to respond by preselecting a default amplitude and direction. In our preceding study, where flexion and extension target ranges were symmetrical, subjects prepared a single default amplitude near the center of each of the two ranges and selected a default direction arbitrarily (Favilla, Hening et al. 1989). Following target presentation, amplitude and direction were specified gradually and in parallel. By using an experimental paradigm in which target amplitudes had unequal probabilities in only one of the two directions, we no sought to determine to what extent the default amplitude prepared by subjects is dependent on the default direction that is selected for a given trial. Five normal subjects produced isometric elbow flexion and extension force impulses to match four targets requiring responses of two amplitudes in each direction. Flexion and extension targets were of the same size. In all biased probability conditions, targets were presented in unpredictable order and, in one direction, the probability of occurrence of the targets was biased (80% versus 20%). In the other direction, the two targets were equiprobable. To maximize the effect of the default settings on response parameters, we required subjects to initiate their response in synchrony with a predictable tone occuring less than a reaction time subsequent to the presentation of the visual target. Such a short interval was chosen to disclose the default parameters selected by limiting the influence of the target just presented on the amplitude and direction of the subjects' responses.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult

Trajectory control in targeted force impulses. VI. Independent specification of response amplitude and direction.

The preceding study of this series (Hening, Favilla and Ghez 1988) examined the time course of the processes by which human subjects use information from a target to set the amplitude of an impulse of isometric elbow force. In that study, subjects were provided with separate cues to time response initiation and to inform them of the required amplitude of the response. When the time between target presentation and response initiation was too brief for them to incorporate information from the target, subjects produced default responses whose amplitudes reflected their prior experience. At longer latencies, subjects specified response amplitude with a gradual time course, starting earlier and ending later than an average reaction time. The present study now examines how two distinct response features, amplitude and direction, are specified following presentation of a target. We sought to answer three main questions. What are the features of responses that are produced before target information is available? Are direction and amplitude specified serially or in parallel? Does the specification of one response feature interfere with the specification of the other? Six normal subjects were studied. They were trained to initiate impulses of isometric elbow force in synchrony with the last of a predictable series of regular tones. The amplitudes and directions were to match those of visual targets requiring flexions or extensions with one of three amplitudes. The targets were presented at random times (0-400 ms) before the last tone. Target directions and amplitudes were either predictable (simple condition) or unpredictable (choice condition). In the simple condition, response amplitudes and directions were independent of the interval between target presentation and response onset (S-R interval). In the choice condition, both amplitude and direction varied with the S-R interval. At short S-R intervals (less than 100 ms), the direction of the subjects' responses was not related to that of the target. The amplitudes of the responses were near the centers of the two target ranges. With increasing S-R intervals, the proportion of correct direction responses gradually increased. Over the same range of S-R intervals, the amplitudes of both right and wrong direction responses to the different targets separated and converged on their respective target amplitudes. Specification of both direction and amplitude was complete at S-R intervals greater than 300 ms. The time course of amplitude specification in this bidirectional paradigm was prolonged over that in a paradigm where response direction was predictable.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult