Search

Background: We examined the validity and reliability of a short robotic test of upper limb proprioception, the Arm Movement Detection (AMD) test, which yields a ratio-scaled, objective outcome measure to be used for evaluating the impact of sensory deficits on impairments of motor control, motor adaptation and functional recovery in stroke survivors. Methods: Subjects grasped the handle of a horizontal planar robot, with their arm and the robot hidden from view. The robot applied graded force perturbations, which produced small displacements of the handle. The AMD test required subjects to respond verbally to queries regarding whether or not they detected arm motions. Each participant completed ten, 60s trials; in five of the trials, force perturbations were increased in small increments until the participant detected motion while in the others, perturbations were decreased until the participant could no longer detect motion. The mean and standard deviation of the 10 movement detection thresholds were used to compute a Proprioceptive Acuity Score (PAS). Based on the sensitivity and consistency of the estimated thresholds, the PAS quantifies the likelihood that proprioception is intact. Lower PAS scores correspond to higher proprioceptive acuity. Thirty-nine participants completed the AMD test, consisting of 25 neurologically intact control participants (NIC), seven survivors of stroke with intact proprioception in the more affected limb (HSS+P), and seven survivors of stroke with impaired or absent proprioception in the more affected limb (HSS-P). Results: Significant group differences were found, with the NIC and HSS+P groups having lower (i.e., better) PAS scores than the HSS-P group. A subset of the participants completed the AMD test multiple times and the AMD test was found to be reliable across repetitions. Conclusions: The AMD test required less than 15 min to complete and provided an objective, ratio-scaled measure of proprioceptive acuity in the upper limb. In the future, this test could be utilized to evaluate the contributions of sensory deficits to motor recovery following stroke.

We have previously demonstrated that, in preparing themselves to aim voluntary impulses of isometric elbow force to unpredictable targets, subjects selected default values for amplitude and direction according the range of targets that they expected. Once a specific target appeared, subjects specified amplitude and direction through parallel processes. Amplitude was specified continuously from an average or central default; direction was specified stochastically from one of the target directions. Using the same timed response paradigm, we now report three experiments to examine how the time available for processing target information influences trajectory characteristics in two-degree-of-freedom forces and multijoint movements. We first sought to determine whether the specification of force direction could also take the form of a discrete stochastic process in pulses of wrist muscle force, where direction can vary continuously. With four equiprobable targets (two force amplitudes in each of two directions separated by 22 degrees or 90 degrees), amplitude was specified from a central default value for both narrow and wide target separations as a continuous variable. Direction, however, remained specified as a discrete variable for wide target separations. For narrow target separations, the directional distribution of default responses suggested the presence of both discrete and central values. We next examined point-to-point movements in a multijoint planar hand movement task with targets at two distances and two directions but at five directional separations (from 30 degrees to 150 degrees separation). We found that extent was again specified continuously from a central default. Direction was specified discretely from alternative default directions when target separation was wide and continuously from a central default when separation was narrow. The specification of both extent and direction evolved over a 200-ms time period beginning about 100 ms after target presentation. As in elbow force pulses, extent was specified progressively in both correct and wrong direction responses through a progressive improvement in the scaling of acceleration and velocity peaks to the target. On the other hand, movement time and hand path straightness did not change significantly in the course of specification. Thus, the specification of movement time and linearity, global features of the trajectories, are given priority over the specific values of extent and direction. In a third experiment, we varied the distances between unidirectional target pairs and found that movement extent is specified discretely, like direction, when the disparity in distances is large. The implications of these findings for contextual effects on trajectory planning are discussed. The independence of extent and direction specification and the prior setting of response duration and straightness provide critical support for the hypothesis that point-to-point movements are planned vectorially.

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.

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)

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)

The present study examines the influences of target predictability, level of practice and response urgency upon the latency and trajectory of a simple motor response. This response is an impulse of isometric force produced by the index finger and aimed to match a step change in a visual target. As expected (Welford 1980), the responses of naive subjects responding as soon as possible after target presentation were initiated at longer latencies when the target steps were of unpredictable amplitudes (choice condition) than when their amplitudes were all the same (simple condition). This choice effect on response latency diminished progressively with practice and eventually disappeared. The trajectories of urgently produced choice responses, however, differed from those of simple responses, and this difference was not reduced by practice. Choice trajectories were more variable and showed a systematic distortion in scaling: response amplitudes exhibited a central tendency bias, or range effect. When targets were equiprobable, responses were biased towards the middle sized target while responses aimed at targets of unequal probability were biased towards the most probable. This effect was independent of the absolute amplitudes of the responses required and was not associated with deviations from the pulse height control policy (Gordon and Ghez 1987a) that human subjects use to vary the amplitude of force impulses. The distortion in scaling and the increased variability of responses aimed at individual targets were markedly reduced when urgency was relaxed and subjects could respond when ready. Then, both simple and choice responses were proportionally scaled to the target, but choice responses were initiated at longer latencies. The changes in trajectory of urgent responses suggests that their inaccuracy occurs because the subjects initiate their responses before the specification of amplitude is complete. The central tendency bias of such incompletely specified responses suggests further that, prior to target presentation, subjects prepare a default response reflecting their expectations. This default may then be modified by information obtained from the target in a process that lasts longer than a minimal reaction time.

In the preceding study (Gordon and Ghez 1987), we showed that accurately targeted isometric force impulses produced by human subjects are governed by a pulse height control policy. Different peak forces were achieved by modulating the rate of rise of force while force rise time was maintained close to a constant value and independent of peak force. An early measure of the rate of rise of force, peak d2F/dt2, was scaled to the required force (target amplitude) and highly predictive of the peak force achieved. In six subjects examined, peak d2F/dt2 accounted for between 70% and 96% of the total variance in peak force. In the present study, we further examined these targeted responses to determine whether the residual variability not predicted by peak d2F/dt2 could be accounted for by adjustments to the force trajectories which compensated for initial errors in the scaling of the d2F/dt2. A statistical model of the determinants of peak force was tested. This model included two paths by which the target amplitude could independently influence the peak force achieved. The first path was preprogrammed pulse height control. In this path, target amplitude determined the initial rate of rise of force (peak d2F/dt2) which in turn determined the final peak force achieved. The second path was an independent influence of errors in the initial scaling of peak d2F/dt2 on peak force. Multiple regression analysis was performed on trajectory variables within the sets of responses by each subject in each condition to determine whether the second path contributed significantly to explaining the variance in peak force. In each subject and condition, there was a significant independent influence of error in d2F/dt2 on peak force, and the direction of this effect was to decrease the magnitudes of peak force errors. These compensatory adjustments accounted for between 1% and 14% of the total variance in peak force. Further multiple regression analyses revealed that inappropriate scaling of the initial phase of the trajectories was compensated for by shortening or prolonging the force rise time. These trajectory adjustments were in turn implemented by modulation of the timing and magnitude of the contractions in the agonist and antagonist muscles that produced the force trajectories. Because these compensatory adjustments were evident in the EMG pattern at latencies too short to be accounted for by peripheral feedback, we assume that they depend on internal monitoring of the unfolding neural commands. These internal feedback processes act in parallel with the programmed commands, both determining the force trajectory. Language: en

(1) Participation of the motor cortex in initiating muscle contraction in an isometric tracking task was assessed in cats trained to make accurate force adjustments using forelimb muscles, in response to a vibrissal/visual display stimulus. Behavior in the task was characterized by short reaction times. While the task was performed, recordings of single cortical units were made in zones within area 4γ defined by the effects of microstimulation in forelimb muscles and by receptive fields on the forelimb. (2) Two types of receptive fields with different regional distributions were observed. Cells with simple receptive fields (superficial or deep) were seen throughout the area sampled, consisting of the lateral half of the anterior and posterior sigmoid gyri. Cells whose receptive fields had complex features (directional specificity, temporal lability, multiple foci, etc.) were preferentially located in the cortex rostral to the cruciate sulcus. (3) The area of motor cortex rostral to the cruciate sulcus also differed from the area caudal to the cruciate sulcus in the timing of task-related activity. Neurons that were active before response onset (lead cells), and could therefore contribute to response initiation, were preferentially located in the rostral cortex, and, in general, had complex receptive fields. (4) Lead cells were active at a constant latency from the stimulus, rather than being timed to response onset. However, the modulation of their activity was related to both the direction and magnitude of the force response. (5) These results suggest that the pericruciate motor cortex of the cat contains two functional subdivisions: a caudal one concerned with ongoing movement, perhaps under the control of specific sensory inputs from the responding limb, and a rostral one involved in initiating movement. Because behaviorally relevant stimuli can rapidly activate a specialized population of cells in the rostral cortex, this area is able to participate in responses with short reaction times.

In a previous study in the cat, we have reported that motor cortex neurons discharging before the initiation of an aimed forearm response (lead cells) are better timed to movement of a display (stimulus) than to the response. The present study was done to distinguish the coding of stimulus and response features in the discharge patterns of such early activity in motor cortex. Single neurons were recorded in the arm area of motor cortex in three cats performing the same pair of responses (forearm flexion and extension) but to display movements in either of the two directions by changing display polarity. The modulation of lead cell activity was contingent on the occurrence of the learned motor response and timed to the stimulus in all conditions. The majority of lead cells (88%, n = 50) fell into one of two distinct classes. In one class of neurons, force-direction (56%, n = 32), activity was contingent on a single direction of forelimb response (flexion or extension) and was thus independent of the direction of the display stimulus. The only muscles whose patterns matched the activity of this class of response-related neurons were forelimb flexors and extensors. In these neurons, the onset of modulation was timed to one or the other of the two stimuli according to the stimulus direction which elicited the appropriate response. Thus, the display-related input to these neurons varied according to the response required. In the second class of neurons, stimulus-direction (32%, n = 18), modulation was associated with a specific stimulus direction rather than the response direction. The pattern of activity of these neurons was similar to the pattern of EMG signals of shoulder and neck muscles during the different task conditions. The contraction of proximal and axial muscles corresponded to a second response elicited by the stimulus, namely attempts at head rotation towards the moving display and was independent of the conditioned forelimb response in both time of onset and direction. To test the possibility that stimulus-direction neurons participated in the control of head rotation we trained two of the animals to also produce isometric changes in neck torque in the direction of the moving display without making the forelimb response. The activity of stimulus-direction neurons was similarly modulated during performance of the neck task. By contrast, force-direction neurons examined during the neck task were either unmodulated or discharged after the neck response.(ABSTRACT TRUNCATED AT 400 WORDS)

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)