Context influences on the preparation and execution of reaching movements

The ability of rapidly adapting our motor behaviour in order to face the unpredictable changes in the surrounding environment is fundamental for survival. To achieve such a high level of efficiency our motor system has to assess continuously the context in which it acts, gathering all available information that can be relevant for planning goal-oriented movements. One still-debated aspect of movement organization is the nature and timing of motor planning. While motor plans are often taken to be concerned with the setting of kinematic parameters as a function of perceptual and motor factors, it has been suggested that higher level, cognitive factors may also affect planning. To explore this issue further, we asked 18 right-handed human participants to perform speeded hand-reaching movement toward a visual target in two different experimental settings, a reaction time (RT) paradigm (go-only task) and a countermanding paradigm. In both tasks participants executed the same movements, but in the countermanding task no-stop trials were randomly intermixed with stop trials. In stop trials participants were required to withhold the ongoing movement whenever a stop signal was shown. It is known that the presence of stop trials induces a consistent increase of the RTs of no-stop trials with respect to the RTs of go-only trials. However, nothing is known about a similar effect for movement times (MTs). We found that RTs and MTs exhibit opposing tendencies, so that a decrease in the RT correspond to an increase in the MT and vice versa. This tendency was present in all our participants and significant in 90% of them. Furthermore we found a moderate, but again very consistent, anticorrelation between RTs and MTs on a trial-by-trial base. These findings are consistent with strategic changes in movement programmes for the very same movements under different cognitive contexts, requiring different degrees of feedback-driven control during movement.     

Inhibitory control of reaching movements in humans

Behavioral flexibility provides a very large repertoire of actions and strategies, however, it carries a cost: a potential interference between different options. The voluntary control of behavior starts exactly with the ability of deciding between alternatives. Certainly inhibition plays a key role in this process. Here we examined the inhibitory control of reaching arm movements with the countermanding paradigm. Right-handed human subjects were asked to perform speeded reaching movements toward a visual target appearing either on the same or opposite side of the reaching arm (no-stop trials), but to withhold the commanded movement whenever an infrequent stop signal was presented (stop trials). As the delay between go and stop signals increased, subjects increasingly failed to inhibit the movement. From this inhibitory function and the reaction times of movements in no-stop trials, we estimated the otherwise unobservable duration of the stopping process, the stop signal reaction time (SSRT). We found that the SSRT for reaching movements was, on average, 206 ms and that it varied with the reaching arm and the target position even though the stop signal was a central stimulus. In fact, subjects were always faster to withhold reaching movements toward visual targets appearing on the same side of the reaching arm. This behavior strictly parallels the course of the reaction times of no-stop trials. These data show that the stop and go processes interacting in this countermanding task are independent, but most likely influenced by a common factor when under the control of the same hemisphere. In addition, we show that the point beyond which the response cannot be inhibited, the so-called point-of-no-return that divides controlled and ballistic phases of movement processing, lies after the inter-hemispheric transfer.     

Impairment of movement initiation and execution but not preparation in idiopathic dystonia

Imaging studies have reported impaired activation of the striatum and their frontal projection sites in dsytonia, areas which are considered to play a role in motor preparation, movement initiation and execution. The aim of this study was to investigate the processes of motor preparation, response initiation and execution in patients with idiopathic torsion dystonia (ITD). We assessed 12 patients with ITD and 12 age-matched controls on a number of reaction time (RT) tasks that differed in degree of motor preparation possible. Subjects performed a visual simple RT (SRT) task, an uncued four-choice reaction time (CRT) task and a fully precued four-choice RT task. A stimulus 1-stimulus 2 (S1-S2) paradigm was used. The warning signal/precue (S1) preceded the imperative stimulus (S2) by either 0 ms (no warning signal or precue) 200 ms, 800 ms, 1,600 ms or 3,200 ms. The patients with ITD had significantly slower RTs and movement times than normals across all RT tasks. The unwarned SRT trials were significantly faster than the uncued CRT trials for both groups. For both groups, precued CRTs were significantly faster than the uncued CRTs. The results show that while response initiation and execution are significantly slower in patients with ITD than normals, movement preparation is not quantitatively or qualitatively different. The results are discussed in relation to previous imaging, behavioural and electrophysiological studies and models of fronto-striatal dysfunction in ITD.     

Distinguishing neural sources of movement preparation and execution. An electrophysiological analysis

The present study examined lateralized event-related potentials (L-ERPs) associated with movement preparation and execution. In a response precuing task that involved hand and foot movements a precue conveyed either information about side and effector, side alone, or no information. Advance movement preparation was indicated by RT shortening with increasing amount of precue information. L-ERPs revealed during the preparatory interval an initial parietal activity when movement side was precued. Later in the preparatory interval L-ERPs revealed a polarity inversion for foot versus hand movements when effector and side were specified in advance. This polarity inversion showed up also in execution-related L-ERP waveforms. Comparison of preparation- versus execution-related brain signals yielded topographic differences and dissimilar dipole sources for hand-related L-ERP activity. We take present findings to indicate that brain generators within the parietal lobe and anterior MI are hierarchically related to precue-induced motor preparation, whilst posterior MI is associated with motor execution functions.     

Influence of cueing on the preparation and execution of untrained and trained complex motor responses

This study investigated the influence of cueing on the performance of untrained and trained complex motor responses. Healthy adults responded to a visual target by performing four sequential movements (complex response) or a single movement (simple response) of their middle finger. A visual cue preceded the target by an interval of 300, 1000, or 2000 ms. In Experiment 1, the complex and simple responses were not previously trained. During the testing session, the complex response pattern varied on a trial-by-trial basis following the indication provided by the visual cue. In Experiment 2, the complex response and the simple response were extensively trained beforehand. During the testing session, the trained complex response pattern was performed in all trials. The latency of the untrained and trained complex responses decreased from the short to the medium and long cue-target intervals. The latency of the complex response was longer than that of the simple response, except in the case of the trained responses and the long cue-target interval. These results suggest that the preparation of untrained complex responses cannot be completed in advance, this being possible, however, for trained complex responses when enough time is available. The duration of the 1st submovement, 1st pause and 2nd submovement of the untrained and the trained complex responses increased from the short to the long cue-target interval, suggesting that there is an increase of online programming of the response possibly related to the degree of certainty about the moment of target appearance.     

Voluntary modification of automatic arm movements evoked by motion of a visual target

We have investigated whether the processes underlying the visually evoked, automatic adjustments to a reach are: (1) modifiable by the subject's intention, and (2) available to initiate movement of a stationary arm. Unpredictable movement of a target (80 m/s through 10 cm, left or right in a third of trials) either evoked a mid-flight adjustment of a reaching movement or else acted as a trigger to start an arm movement. Subjects were instructed to respond as rapidly as possible by moving their finger either in the same or in the opposite direction to the target. The target shift evoked an early (125-160 ms) and/or a later (> 160 ms) class of response in the reaching arm. The early response was highly automatic in that it could not be reversed (move opposite) by the subjects' intention. However, the subjects' intention did influence the frequency of occurrence and the size of this early response. The later response was totally modifiable in that it changed direction according to the subjects' intention. Similar classes of response were observed in stationary limbs, but the early, more automatic response was substantially weaker than that elicited during a reach. Two possible mechanisms are proposed to explain these results. The first is a dual-pathway model, which assumes that the two response classes are each generated by separate visuo-motor processes with different properties. The second model assumes both responses are generated by a single visuo-motor mechanism that is under the control of a higher, attentional process.     

Motor imagery influences the execution of repetitive finger opposition movements

Motor imagery (MI) is the ability to imagine performing a movement without executing it. In literature, there have been numerous reports on the influence of MI on motor practice and the beneficial effects of “mental practice” on the physical performance has been suggested to rely to the close temporal association between motor rehearsal and actual performance. In the present study, we aimed to evaluate whether the addition of a period of motor imagery between two motor practice trials could modify movement execution in a repetitive finger opposition motor task performed at maximal speed and whether the effect of motor imagery on motor practice is dependant on the complexity of movement. We observed that the addition of motor imagery to the sole motor practice was able to influence the performance of repetitive finger opposition movements inducing an increase of the velocity of movement greater than that observed with the motor practice alone. Further the addition of motor imagery was able to induce a modification in the motor strategy in terms of duration of the main phases of movements. This was more evident when subjects executed a finger sequential task with respect to a simple finger tapping task. We assume that mental rehearsal facilitates the brain network involved in sensorimotor control, particularly acting on those neural structures involved in the motor program.     

Colour vision can contribute to fast corrections of arm movements

Can chromatic information be used for the fast on-line control of action? In order to find out we asked subjects to tap a red square as quickly as possible. In half of the trials the red square’s position changed as soon as the subject’s hand started to move. We examined how quickly subjects could adjust their movements to this change. In half of the trials there was also a green square of the same luminance as the red one. If there was a green square, and the red square’s position changed, the change consisted of the two squares exchanging places, so that all that really changed was the squares’ colours. In such cases subjects could not have adjusted their movements without having analysed the colour. Nevertheless, subjects could respond adequately within as little as 120 ms. This was even so if the squares’ luminances changed considerably at the moment that the subject’s hand started to move. Thus, chromatic information can be used for the fast on-line control of action     

Motor programmes for goal-directed movements are continuously adjusted according to changes in target location

Summary We have studied fast arm movements in response to double-step stimuli in two-dimensional space. In a previous paper we found that such movements did not start in the direction of the first or the second target, but in a direction between the two targets. The initial movement direction was found to depend in a continuous fashion on the inter-stimulus interval and on the reaction time. Therefore we concluded that the internal representation of a discrete target displacement is a gradually shifting internal target, moving from the first to the second target location. In this paper we investigate whether the arm movements also show a modification of the trajectory during the movement. An inter-stimulus interval of 100 ms was chosen, because then the initial movement direction is the same as in the response to a single-step displacement. We found that on average double-step trajectories deviate significantly from their original trajectory within 60 ms, and in some cases even within 30 ms of the start of the movement. We conclude that a motor programme is centrally modified according to a changed target location. We hypothesize that the generation of the motor programme starts after the target presentation, and that the activation levels for the appropriate muscles are continuously adjusted to move the hand in the direction of the current internal representation of the target.     

Stability properties of human reaching movements

Through an experimental study of the stability properties of the human neuromuscular system while it performs simple point-to-point arm movements, this paper evaluates the concepts of equilibrium and virtual trajectories as a means of executing movement of the arm. Human subjects grasped the instrumented handle of a two-link robot manipulandum and performed specified point-to-point planar arm trajectories. Computer-controlled brakes were used to subtly change the movements by constraining the trajectory along an arc of radius equal to the length of one link of the manipulandum. Target points were arranged to lie along the arc so that the subject could complete the movement even when constrained. Three situations were tested: (1) unconstrained throughout the movement, (2) constrained through the entire movement, and (3) initially constrained and then released during movement. Experimental results showed that the constraint evoked significant forces strongly oriented so as to restore the hand to the unconstrained hand path. In addition, when released from the constraint, these forces caused a strong tendency to return the hand to the unconstrained path before the end of the movement was reached. Such strong positional stability properties of the arm reinforce the notion that a moving attractor point dominates the dynamics of the arm during movement. Additionally, bounds on the shape of the virtual trajectory were found which indicate that the equilibrium point remains close to the actual movement produced. These results, showing that a controlled equilibrium point may be used for planning and coordinating multijoint movements, are consistent with an equilibrium point hypothesis.     

Proprioceptive guidance and motor planning of reaching movements to unseen targets

The ability to make accurate reaching movements toward proprioceptively defined target locations was studied in seven normal subjects who were trained to reach to five different targets in a horizontal plane, with no vision of hand or target. The task consisted of moving a handle from a fixed origin to each target location, fast and accurately. Target locations were learned in training sessions that utilized acoustic cuing. Most movements were rapid, with a bell-shaped velocity profile. The error in target reproduction, which constituted the difference between the position consciously identified as the correct target location and the real target location, was calculated in each trial. This was compared with the error in preprogrammed reaching, which constituted the difference between the point in space where the initial fast movement toward the target ended and the target location. The absence of significant differences between these two error types indicated that the transformation from an internal representation of target location into a motor program for reaching to it did not introduce an additional reaching error. Learning of target locations was done only with the right hand, yet, reaching of both hands was tested. This allowed a comparison between the subjects' ability to utilize a transformed spatial code (reaching with the untrained hand) and their ability to use a direct sensory-motor code (reaching with the trained hand). While transformation of the spatial code was found to reduce it's accuracy, utilization of this code in motor programming again did not appear to introduce an additional error.     

Perceptual distortion contributes to the curvature of human reaching movements

Unconstrained point-to-point human arm movements are generally gently curved, a fact which has been used to assess the validity of models of trajectory formation. In this study we examined the relationship between curvature perception and movement curvature for planar sagittal and transverse arm movements. We found a significant correlation (P<0.0001, n=16) between the curvature perceived as straight and the curvature of actual arm movements. We suggest that subjects try to make straight-line movements, but that actual movements are curved because visual perceptual distortion makes the movements appear to be straighter than they really are. We conclude that perceptual distortion of curvature contributes to the curvature seen in human point-to-point arm movements and that this must be taken into account in the assessment of models of trajectory formation.     

Effect of load level and muscle pain intensity on the motor control of elbow-flexion movements

This study assessed interactions between mild/moderate muscle pain and inertial load on the control of human elbow-flexion movements. It is hypothesized that high inertial load combined with moderate muscle pain intensity affect the motor control more than for low inertial-load combined with low-intensity pain. Fifteen subjects performed horizontal pointing movements (70° range) under three load conditions: 0, 4, and 10 kg. Pain was induced by injection of 0.5 ml and 1.5 ml hypertonic saline into the biceps muscle. Subjects scored the muscle pain intensity on a visual analogue scale (VAS). Elbow joint position, VAS, and the electromyograms (EMG, m. biceps brachii, m. triceps brachii, m. brachioradialis, and m. trapezius) were recorded. Mild and moderate muscle pain attenuated acceleration profiles [6.1(0.9)%], effective movement amplitude [3.2 (0.7)%], peak velocity [5.8 (0.9)%] and prolonged the reaction time [21 (5)%]. No interaction between muscle pain intensity and inertial load was found for the kinematic parameters. EMG profiles from m. biceps brachii, m. triceps brachii, and m. brachioradialis were similarly attenuated [10.2 (0.80)%] by mild and moderate muscle pain in all inertial load conditions. For high inertial load, the initial agonist EMG burst activity was more attenuated [50 (5.3)%] by moderate muscle pain compared with mild muscle pain [34 (4.2)%]. These data suggest that for high effort-demanding tasks muscle pain differently affects the motor planning according to the pain-intensity level. Perturbations of motor planning lead to changes on movement strategies, which might be a potential cause of musculoskeletal problems.     

Behavioral and electrocortical evidence of an interaction between probability and task metrics in movement preparation

Recent neurophysiological evidence suggests that cognitive factors shape neural activity in cortical areas such as parietal (area 5), premotor, and primary motor cortex. The implication of these findings is that behavioral signatures of cognitive factors and movement-specific factors should likewise be interdependent. The present study provides evidence of this interdependence in both behavioral (reaction time) and electrophysiological (P300) measures. Subjects performed a two-choice pointing task, in which the angular distance between the two required movement directions (task metrics) and the probability of the two responses was varied. In a control condition, a single reaction was required in response to both stimuli to test for the influence of stimulus metrics. Results from the pointing task showed a clear interaction between the metrics and effects of probability. When the potential targets were widely separated (120 degrees), stimulus probability influences reaction time and P300 amplitude in the classic fashion (longer reaction times and larger P300 amplitudes to less probable responses). When pointing to targets that were narrowly separated (20 degrees), probability had no effect: both rare and frequent targets were "functionally frequent." The same interaction was not observed in the control condition, indicating that metrics were primarily influencing movement preparation rather than stimulus processing. The results are consistent with the theoretical framework of dynamic field theory and demonstrate that metrics are an important factor that must be taken into account when assessing the processes associated with movement preparation.     

Anticipatory postural adjustments during self-paced and reaction-time movements

We studied the changes in the anticipatory postural adjustments (APAs), associated with dropping a load from extended arms and during fast bilateral shoulder flexion movements, when movements were performed in a self-paced manner and under a simple reaction-time instruction. The latter instruction applied time pressure and did not allow the regular pattern of APAs to be used. In particular, the following questions were asked: (1) are there changes in the relative timing of APAs under the reaction time condition; (2) are changes in the relative timing of APAs associated with changes in APAs themselves; (3) can different postural strategies be used to maintain stability under self-paced and reaction time conditions; and (4) are changes in APAs related to actual reaction time or to a change in the instruction? In particular, under reaction-time conditions, APAs occurred later in time, typically simultaneously with the initiation of the focal movement. Additional changes in electromyographic (EMG) patterns in postural muscles included an increase in the amplitude of EMG bursts and “speeding-up” some of the tri-phasic patterns in postural dorsal-ventral muscle pairs. This was accompanied by a smaller early shift of the center of pressure followed by its more rapid delayed displacement. There was considerable variability in the changes of EMG and dynamic characteristics across subjects. Some of the changes in the EMG patterns in postural muscles depended on actual reaction time, while others were related to a change in the instruction and occurred even if actual reaction times were long enough to allow for the typical self-paced APA patterns to occur. These findings can be interpreted as supporting the parallel control hypothesis for the focal movement and postural adjustments. Alternatively, they can be interpreted within a framework that implies the generation of a single control function, which is transformed into two components, one directed at the focal muscles/joints and the other directed at postural muscles/joints.     

The effects of changing movement velocity and complexity on response preparation: evidence from latency,kinematic, and EMG measures

Summary If movement control is afforded through the advance planning, or preprogramming, of upcoming actions, then one of the behavioral outcomes should be an increase in reaction time (RT) as the movement becomes more complex. In some situations, however, RT does not increase across levels of complexity, rather it remains invariant. In these cases, on-line preparation is typically inferred. That is, the sequence is said to be prepared in parts throughout the movement, as opposed to entirely beforehand. Given that there is some planning occurring during the sequence, then evidence of this process should be apparent within the movement itself. Three such dependent variables appear to provide such evidence. Specifically, the number of times the underlying accelerations cross the zero line within the movement, the number of significant deviations within the acceleration trace, and the length of time for which the muscles are active (as measured by EMG) in relation to the duration of the movement. In the present experiment, then, these variables were measured in addition to the time required to prepare and initiate a movement performed under conditions conducive to either preprogramming or online preparation. Specifically, the movements were either completed as fast as possible, or at a considerably slower, more controlled speed. Each of the dependent variables displayed evidence of preprogramming in the movements completed at the fast velocity, and on-line preparation in the slower paced movements. Thus, in the slow condition, subjects appeared to rely more heavily on on-line prepared adjustments to produce an accurate outcome. The convergence attained between the various dependent measures lends power to the conclusions regarding hypothesized modes of control within the different speeds of movement.     

Discrete and continuous planning of hand movements and isometric force trajectories

 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° or 90°), 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° to 150° 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. Received: 6 August 1996 / Accepted: 18 December 1996     

Movement preparation and working memory: a behavioural dissociation

We can cross temporal sensorimotor contingencies by remembering sensory events or by anticipating motor responses. Here we tested the hypothesis that sensory and motor representations can be accessed according to different temporal dynamics. We predicted that the manipulation of movement representations would lead to a performance independent from the length of a delay interposed between sensory instructions and behavioural responses. Conversely, we expected a delay-dependent performance whenever temporary storage of sensory information was necessary to solve the task. We have measured reaction times and error rate in subjects performing a delayed non-matching to sample task. Task contingencies rather than explicit instructions ensured that either sensory or motor representations were used to cross the delay period on each trial. We tested our hypothesis by manipulating the length of the delays between stimulus presentation and behavioural response. We found that carrying sensory material over temporal gaps affects performance as a non-linear function of time, whereas movement representations remain robust over a wide range of delays. This novel behavioural paradigm might prove effective in dissociating the neural bases of preparatory and mnestic processes in normal human subjects, as well as their disorders in neurological patients.     

Motor preparation in a memorised delay task

The effect on reaction time (RT) and movement time (MT) of remembering which one of several targets to move to was investigated in 18 participants who completed 416 trials in each task. On each trial, participants moved their index finger from a central, illuminated switch (the stimulus) to one of eight targets located on the circumference of a 6 cm radius circle. A visual cue (illumination of the target) informed the participant of the appropriate target. In the memorised delay task, the cued target was lit for 300 ms followed by a variable (450–750 ms) foreperiod during which the participant was required to remember the location of the target until the stimulus light was extinguished. In the non-memorised delay task, the target remained lit during the entire foreperiod (750–1050 ms) until the response was completed. At the go signal (stimulus light extinguished) participants moved as quickly and accurately as possible to the cued target. Both RT and MT were significantly (p<0.05) longer in the memorised delay task. The increase in RT shows that remembering which target imposed a greater load on motor preparation even though all the information needed for preparing the response was presented in the cue at the beginning of each trial. The increase in MT raises the possibility that movement execution was also programmed during motor preparation.     

Differential relation of discharge in primary motor cortex and premotor cortex to movements versus actively maintained postures during a reaching task

The activity of cells in primary motor cortex (MI) and dorsal premotor cortex (PMd) were compared during reaching movements in a reaction-time (RT) task, without prior instructions, which required precise control of limb posture before and after movement. MI neurons typically showed strong, directionally tuned activity prior to and during movement as well as large gradations of tonic activity while holding the limb over different targets. The directionality of their movementand posture-related activity was generally similar. Proximal-arm muscles behaved similarly. This is consistent with a role for MI in the moment-to-moment control of motor output, including both movement and actively maintained postures, and suggests a common functional relation for MI cells to both aspects of motor behavior. In contrast, PMd cells were generally more phasic, frequently emitting only strong bursts of activity confined mainly to the behavioral reaction time before movement onset. PMd tonic activity during different postures was generally weaker than in MI, and showed a much more variable relation with their movement-related directional tuning. These results imply that the major contribution of PMd to this RT task occurred prior to the onset of movement itself, consistent with a role for PMd in the selection and planning of visually guided movements. Furthermore, the nature of the relative contribution of PMd to movement versus actively maintained postures appears to be fundamentally different from that in MI. Finally, there was a continuous gradient of changes in responses across the rostrocaudal extent of the precentral gyrus, with no abrupt transition in response properties between PMd and MI.