Development of Force Adaptation During Childhood

Humans learn to make reaching movements in novel dynamic environments by acquiring an internal motor model of their limb dynamics. Here, the authors investigated how 4- to 11-year-old children (N = 39) and adults (N = 7) adapted to changes in arm dynamics, and they examined whether those data support the view that the human brain acquires inverse dynamics models (IDM) during development. While external damping forces were applied, the children learned to perform goal-directed forearm flexion movements. After changes in damping, all children showed kinematic aftereffects indicative of a neural controller that still attempted to compensate the no longer existing damping force. With increasing age, the number of trials toward complete adaptation decreased. When damping was present, forearm paths were most perturbed and most variable in the youngest children but were improved in the older children. The findings indicate that the neural representations of limb dynamics are less precise in children and less stable in time than those of adults. Such controller instability might be a primary cause of the high kinematic variability observed in many motor tasks during childhood. Finally, the young children were not able to update those models at the same rate as the older children, who, in turn, adapted more slowly than adults. In conclusion, the ability to adapt to unknown forces is a developmental achievement. The present results are consistent with the view that the acquisition and modification of internal models of the limb dynamics form the basis of that adaptive process.     

Development of force adaptation during childhood

Humans learn to make reaching movements in novel dynamic environments by acquiring an internal motor model of their limb dynamics. Here, the authors investigated how 4- to 11-year-old children (N = 39) and adults (N = 7) adapted to changes in arm dynamics, and they examined whether those data support the view that the human brain acquires inverse dynamics models (IDM) during development. While external damping forces were applied, the children learned to perform goal-directed forearm flexion movements. After changes in damping, all children showed kinematic aftereffects indicative of a neural controller that still attempted to compensate the no longer existing damping force. With increasing age, the number of trials toward complete adaptation decreased. When damping was present, forearm paths were most perturbed and most variable in the youngest children but were improved in the older children. The findings indicate that the neural representations of limb dynamics are less precise in children and less stable in time than those of adults. Such controller instability might be a primary cause of the high kinematic variability observed in many motor tasks during childhood. Finally, the young children were not able to update those models at the same rate as the older children, who, in turn, adapted more slowly than adults. In conclusion, the ability to adapt to unknown forces is a developmental achievement. The present results are consistent with the view that the acquisition and modification of internal models of the limb dynamics form the basis of that adaptive process.     

Stiffness control after fast goal-directed arm movements

Mechanical parameters of the effector system directly after the termination of fast goal-directed arm movements were studied.Subjects were asked to move their hand as fast as possible to a target the instant the target was presented. Only movements of the subjects' forearms were allowed. They were also instructed not to react actively to forces applied suddenly to their forearm after the movement. As a result of such a force pulse the arm moved to a new position. The apparent stiffness, i.e. the quotient of the applied force and the resultant change of position, was measured. This stiffness is a measure for the resistance of the forearm to externally applied mechanical disturbances.It was found that after the arm has reached the target the apparent stiffness decreases as a function of time. This is an agreement with the declining amplitude of the electromyographic activity of the muscles that effect the movement.Arguments are given to support the hypothesis that this apparent stiffness control is part of the motor programme for movements of the forearm, i.e. the stiffness is planned together with the movement.     

Adaptive motor behavior of cerebellar patients during exposure to unfamiliar external forces

The authors investigated adaptation of goal-directed forearm movements to an unknown external viscous force assisting forearm flexion in 6 patients with cerebellar dysfunction and in 6 control participants. Motor performance was generally degraded in cerebellar patients and was markedly reduced under the force condition in both groups. However, patients and controls were able to adapt to the novel force within 8 trials. Only the healthy controls were able to improve motor performance when readapting to a null-force condition. The results indicate that cerebellar patients' motor control system has imprecise estimations of actual limb dynamics at its disposal. Force adaptation may have been preserved because single-joint movements were performed, whereas the negative viscous force alone and no interaction forces had to be compensated.     

Motor skill learning in the middle-aged: limited development of motor chunks and explicit sequence knowledge

The present study examined whether middle-aged participants, like young adults, learn movement patterns by preparing and executing integrated sequence representations (i.e., motor chunks) that eliminate the need for external guidance of individual movements. Twenty-four middle-aged participants (aged 55–62) practiced two fixed key press sequences, one including three and one including six key presses in the discrete sequence production task. Their performance was compared with that of 24 young adults (aged 18–28). In the middle-aged participants motor chunks as well as explicit sequence knowledge appeared to be less developed than in the young adults. This held especially with respect to the unstructured 6-key sequences in which most middle-aged did not develop independence of the key-specific stimuli and learning seems to have been based on associative learning. These results are in line with the notion that sequence learning involves several mechanisms and that aging affects the relative contribution of these mechanisms.     

Vector coding in slow goal-directed arm movements

It has been found that the estimate of relative target direction is consistently biased. Relative target direction refers to the direction in which a target is located relative to another location in space (e.g., a starting position in the case of goal-directed movements). In this study, we have tested two models that could underlie this biased estimate. The first proposed model is based on a distorted internal representation of locations (i.e., we perceive a target at the “wrong” location). We call this thedistorted location model. The second model is based on the idea that the derivation of target direction from spatial information about starting and target position is biased. We call this thebiased direction model. These two models lead to different predictions of the deviations that occur when the distance between the starting position and the target position is increased. Since we know from previous studies that the initial direction of slow arm movements reflects the target direction estimate, we tested the two models by analyzing the initial direction of slow arm movements. The results show that the biased direction model can account for the biases we find in the target direction estimate for various target distances, whereas the distorted location model cannot. In two additional experiments, we explored this model further. The results show that the biases depend only on the orientation of the line through starting position and target position relative to the plane through longitudinal head or body axis and starting position. We conclude that the initial part of (slow) goal-directed arm movements is planned on the basis of a (biased) target direction estimate and not on the basis of a wrong internal representation of target location. This supports the hypothesis that we code displacements of our limbs in space as a vector.     

Baby steps: investigating the development of perceptual–motor couplings in infancy

There are cells in our motor cortex that fire both when we perform and when we observe similar actions. It has been suggested that these perceptual‐motor couplings in the brain develop through associative learning during correlated sensorimotor experience. Although studies with adult participants have provided support for this hypothesis, there is no direct evidence that associative learning also underlies the initial formation of perceptual–motor couplings in the developing brain. With the present study we addressed this question by manipulating infants’ opportunities to associate the visual and motor representation of a novel action, and by investigating how this influenced their sensorimotor cortex activation when they observed this action performed by others. Pre‐walking 7–9‐month‐old infants performed stepping movements on an infant treadmill while they either observed their own real‐time leg movements (Contingent group) or the previously recorded leg movements of another infant (Non‐contingent control group). Infants in a second control group did not perform any steps and only received visual experience with the stepping actions. Before and after the training period we measured infants’ sensorimotor alpha suppression, as an index of sensorimotor cortex activation, while they watched videos of other infants’ stepping actions. While we did not find greater sensorimotor alpha suppression following training in the Contingent group as a whole, we nevertheless found that the strength of the visuomotor contingency experienced during training predicted the amount of sensorimotor alpha suppression at post‐test in this group. We did not find any effects of motor experience alone. These results suggest that the development of perceptual–motor couplings in the infant brain is likely to be supported by associative learning during correlated visuomotor experience.     

Directing attention to movement effects enhances learning: a review

Studies investigating the influence of the learner’s focus of attention, induced by instructions or feedback, on motor skill learning are reviewed. In general, directing performers’ attention to the effects of their movements (external focus of attention) appears to be more beneficial than directing their attention to their own movements (internal focus of attention). Preliminary evidence is presented indicating that an internal attentional focus constrains the motor system by interfering with natural control processes, whereas an external focus seems to allow automatic control processes to regulate the movements. Support for the view that actions are controlled by their anticipated effects comes from research demonstrating functional variability in motor control, as well as the benefits of purposeful activity in occupational therapy. We explain these results in terms of the ideomotor principle of human actions (James, 1890) and its more modern derivatives (Hommel, 1996; Prinz, 1990, 1997).     

A study of a bimanual synergy associated with holding an object

The task of supporting an object with one or two hands was used to test the applicability of the notion of synergy. Subjects sat with their dominant forearm supported up to the wrist while holding a cylindrical “cup” between their thumb and fingers. Force transducers recorded the grip force applied normal to the cup's side by the thumb and the force applied normal to the cup bottom. On different series, a supporting force was added to and released from the bottom of the cup by the subject's non-dominant hand or by the experimenter. As predicted, the results indicated feedforward adjustments of the grip force, and of the EMGs, and significant correlations between grip force and supporting force when they were produced by two hands of one person, and the lack of such closely tied changes when the two forces were produced by two different persons. In the latter case, different subjects could demonstrate grip force changes in different directions. The findings suggest that grip force adjustments represented peripheral patterns of a single central process (a single synergy) rather than being separately controlled focal and postural components of the action.PsycINFO classification: 2330     

Changes in pinch force with bidirectional load forces

The forces used to grasp an object were measured while positive (push) and negative (pull) load forces were applied to the hand under varying frictional conditions. Subjects held between the tips of their thumb and index finger a manipulandum composed of two symmetrically mounted disks. The manipulandum was connected to the stage of an electromagnetic linear motor that generated load forces under computer control. In the first experiment, subjects held the position of the manipulandum constant while the motor generated forces in first the positive and then the negative direction. The motor force at which the manipulandum slipped from the fingers was measured in the second experiment. In both experiments, friction was varied by changing the surface (sandpaper, suede, or plastic) of the manipulandum disks. The pinch forces produced by subjects were linearly related to changes in motor force in both the positive and negative directions, with the slope of this relation varying as a function of the surface properties of the manipulandum. The modulation of pinch force with motor force was influenced, however, by the direction of the load force; higher forces were produced in response to negative load forces. Slip forces varied as a function of pinch force and surface texture; higher forces were associated with materials with lower coefficients of friction. These findings suggest that the friction between the skin and an object being grasped changes as a function of the direction of force that the object applies to the skin, possibly due to the anisotropic nature of glabrous skin, and that this mechanical property contributes to variations in pinch force.     

Relative damping improves linear mass-spring models of goal-directed movements

A limitation of a simple linear mass-spring model in describing goal directed movements is that it generates rather slow movements when the parameters are kept within a realistic range. Does this imply that the control of fast movements cannot be approximated by a linear system? In servo-control theory, it has been proposed that an optimal controller should control movement velocity in addition to position. Instead of explicitly controlling the velocity, we propose to modify a simple linear mass-spring model. We replaced the damping relative to the environment (absolute damping) with damping with respect to the velocity of the equilibrium point (relative damping). This gives the limb a tendency to move as fast as the equilibrium point. We show that such extremely simple models can generate rapid single-joint movements. The resulting maximal movement velocities were almost equal to those of the equilibrium point, which provides a simple mechanism for the control of movement speed. We further show that peculiar experimental results, such as an 'N-shaped' equilibrium trajectory and the difficulties to measure damping in dynamic conditions, may result from fitting a model with absolute damping where one with relative damping would be more appropriate. Finally, we show that the model with relative damping can be used to model subtle differences between multi-joint interceptions. The model with relative damping fits the data much better than a version of the model with absolute damping.     

Gestures in Instructional Animations: A Helping Hand to Understanding Non‐human Movements?

Recent research on dynamic visualizations suggests that these visualizations are effective for learning human movements such as knot tying or paper folding. Using embodied theories of cognition, this study investigated whether learning non‐human movements from a dynamic visualization can also be enhanced by grounding these movements in the learner's motor system. University students viewed an animation on lightning formation, and followed the animation's movements with gestures, saw an on‐screen human hand follow the movements or saw an arrow follow the movements. Results showed that observing an on‐screen human hand following the movements in the animation, but not actually performing these movements, enhanced retention and transfer performance compared with watching the animation without gestures. This suggests that observation of human hand movement in animation‐based instruction, which plays an important role in learning of procedural‐motor tasks, can also improve people's learning from other dynamic systems. Copyright © 2013 John Wiley & Sons, Ltd.     

Curved saccade trajectories reveal conflicting predictions in associative learning

We report how the trajectories of saccadic eye movements are affected by memory interference acquired during associative learning. Human participants learned to perform saccadic choice responses based on the presentation of arbitrary central cues A, B, AC, BC, AX, BY, X, and Y that were trained to predict the appearance of a peripheral target stimulus at 1 of 3 possible locations, right (R), mid (M), or left (L), in the upper hemifield. We analyzed as measures of associative learning the frequency, latency, and curvature of saccades elicited by the cues and directed at the trained locations in anticipation of the targets. Participants were trained on two concurrent discrimination problems A+R, AC+R, AX+M, X+M and B+L, BC+L, BY+M, Y+M. From a connectionist perspective, cues were predicted to acquire associative links connecting the cues to the trained outcomes in memory. Model simulations based on the learning rule of the Rescorla and Wagner (1972) model revealed that for some cues, the prediction of the correct target location was challenged by the interfering prediction of an incorrect location. We observed that saccades directed at the correct location in anticipation of the target curved away from the location that was predicted by the interfering association. Furthermore, changes in curvature during training corresponded to predicted changes in associative memory. We propose that this curvature was caused by the inhibition of the incorrect prediction, as previously has been suggested with the concept of distractor inhibition (Sheliga, Riggio, & Rizzolatti, 1994; Tipper, Howard, & Houghton, 2000). The paradigm provides a new method to examine memory interference during associative learning.     

Visuomotor Adaptation and Proprioceptive Recalibration

ABSTRACT

Motor learning, in particular motor adaptation, is driven by information from multiple senses. For example, when arm control is faulty, vision, touch, and proprioception can all report on the arm's movements and help guide the adjustments necessary for correcting motor error. In recent years we have learned a lot about how the brain integrates information from multiple senses for the purpose of perception. However, less is known about how multisensory data guide motor learning. Most models of, and studies on, motor learning focus almost exclusively on the ensuing changes in motor performance without exploring the implications on sensory plasticity. Nor do they consider how discrepancies in sensory information (e.g., vision and proprioception) related to hand position may affect motor learning. Here, we discuss research from our lab and others that shows how motor learning paradigms affect proprioceptive estimates of hand position, and how even the mere discrepancy between visual and proprioceptive feedback can affect learning and plasticity. Our results suggest that sensorimotor learning mechanisms do not exclusively rely on motor plasticity and motor memory, and that sensory plasticity, in particular proprioceptive recalibration, plays a unique and important role in motor learning.     

Rhesus monkeys (<Emphasis Type="Italic">Macaca mulatta</Emphasis>) show robust evidence for memory awareness across multiple generalization tests

The possibility that memory awareness occurs in nonhuman animals has been evaluated by providing opportunity to decline memory tests. Current evidence suggests that rhesus monkeys (Macaca mulatta) selectively decline tests when memory is weak (Hampton in Proc Natl Acad Sci USA 98:5359–5362, 2001; Smith et al. in Behav Brain Sci 26:317–374, 2003). However, much of the existing research in nonhuman metacognition is subject to the criticism that, after considerable training on one test type, subjects learn to decline difficult trials based on associative learning of external test-specific contingencies rather than by evaluating the private status of memory or other cognitive states. We evaluated whether such test-specific associations could account for performance by presenting monkeys with a series of generalization tests across which no single association with external stimuli was likely to adaptively control use of the decline response. Six monkeys performed a four alternative delayed matching to location task and were significantly more accurate on trials with a decline option available than on trials without it, indicating that subjects selectively declined tests when memory was weak. Monkeys transferred appropriate use of the decline response under three conditions that assessed generalization: two tests that weakened memory and one test that enhanced memory in a novel way. Bidirectional generalization indicates that use of the decline response by monkeys is not controlled by specific external stimuli but is rather a flexible behavior based on a private assessment of memory.     

Changes in Movement Symmetry Associated With Strengthening and Fatigue of Agonist and Antagonist Muscles

The hypothesis that strengthening or fatiguing procedures applied on active muscles can affect the symmetry of rapid, discrete movements was tested. Subjects (N = 12) performed rapid, consecutive elbow flexions and extensions between 2 targets before and after (a) applying a strength training program, (b) fatiguing elbow flexors, and (c) fatiguing elbow extensors. The results demonstrated that an increase in strength of elbow extensors caused by applied strength training is associated with an increase in the symmetry ratio (i.e., acceleration time divided by deceleration time) of elbow flexion movements. The symmetry ratio also increased and decreased in movements when agonists and antagonists were fatigued, respectively. Because the strength training and fatiguing procedures are both known to affect muscle force, the data are interpreted as changes in muscles' ability to exert the force while acting as agonists or antagonists. Namely, muscles need equal impulses of force (torque multiplied by time) to accelerate and, thereafter, to decelerate the limb while performing a rapid, discrete movement. The symmetry ratio may therefore be changed so that more time will be provided for muscles that become relatively weaker (compared with their antagonists) because a strengthening or fatiguing procedure has been applied, whereas a shorter time period should be sufficient for action of their stronger antagonists. Although, in the literature, the studied phenomenon has been discussed as a predominantly motor control phenomenon, the present data suggest that the movement symmetry could also be related to agonists' and antagonists' ability to exert force, particularly while performing rapid, discrete movements.     

Atypical inter‐hemispheric communication correlates with altered motor inhibition during learning of a new bimanual coordination pattern in developmental coordination disorder

Impairment of motor learning skills in developmental coordination disorder (DCD) has been reported in several studies. Some hypotheses on neural mechanisms of motor learning deficits in DCD have emerged but, to date, brain‐imaging investigations are scarce. The aim of the present study is to assess possible changes in communication between brain areas during practice of a new bimanual coordination task in teenagers with DCD (n = 10) compared to matched controls (n = 10). Accuracy, stability and number of mirror movements were computed as behavioural variables. Neural variables were assessed by electroencephalographic coherence analyses of intra‐hemispheric and inter‐hemispheric fronto‐central electrodes. In both groups, accuracy of the new coordination increased concomitantly with right intra‐hemispheric fronto‐central coherence. Compared to typically developing teenagers, DCD teenagers presented learning difficulties expressed by less stability, no stabilization of the new coordination and a greater number of mirror movements despite practice. These measures correlated with reduced inter‐hemispheric communication, even after practice of the new coordination. For the first time, these findings provide neuro‐imaging evidence of a kind of inter‐hemispheric ‘disconnection’ related to altered inhibition of mirror movements during motor learning in DCD.     

Autonomy facilitates repeated maximum force productions

Performer autonomy (or self-control) has consistently been shown to enhance motor learning, and it can also provide immediate benefits for motor performance. Autonomy is also a key variable in the OPTIMAL theory of motor learning (Wulf & Lewthwaite, 2016). It is assumed to contribute to enhanced expectancies and goal-action coupling, affecting performance effectiveness and efficiency. The purpose of the present study was to examine whether providing autonomy support by giving performers choices would enhance their ability to maintain maximum force levels. Participants were asked to repeatedly produce maximum forces using a hand dynamometer. After 2 initial trials with the dominant and non-dominant hand, stratified randomization was used to assign participants with the same average maximum force to one of two groups, choice or yoked control groups. Choice group participants were able to choose the order of hands (dominant, non-dominant) on the remaining trials (3 per hand). For control group participants, hand order was determined by choice-group counterparts. Maximum forces decreased significantly across trials in the control group, whereas choice group participants were able to maintain the maximum forces produced on the first trial. We interpret these findings as evidence that performer autonomy promotes movement efficiency. The results are in line with the view that autonomy facilitates the coupling of goals and actions (Wulf & Lewthwaite, 2016).     

Eye-hand coordination: spatial localization after saccadic and pursuit eye movements

Subjects produced speeded and unspeeded hand movements to a target location after either saccadic or pursuit eye movements to the target. Hand movements began either aligned with the initial position of gaze or from some other location. Subjects generally underestimated the extent of the pursuit eye movements relative to estimates made after saccades. With speeded hand movements, however, the underestimation was reduced considerably if the hand movements began aligned with a location other than the initial position of gaze. The results reveal details of the mechanisms underlying eye-hand coordination and show that important differences exist in the information used for localization for slow and rapid limb movements.     

Characteristics of Unintentional Movements by a Multijoint Effector

The authors explored the phenomenon of unintentional changes in the equilibrium state of a multijoint effector produced by transient changes in the external force. The subjects performed a position-holding task against a constant force produced by a robot and were instructed not to intervene voluntarily with movements produced by changes in the robot force. The robot produced a smooth force increase leading to a hand movement, followed by a dwell time. Then, the force dropped to its initial value leading to hand movement toward the initial position, but the hand stopped short of the initial position. The undershoot magnitude increased linearly with the peak hand displacement and exponentially with dwell time (time constant of about 1 s). For long dwell times, the hand stopped at about half the total distance to the initial position. The authors interpret the results as consequences of a drift of the referent hand coordinate. Our results provide support for back-coupling between the referent and actual body configurations during multijoint actions and produce the first quantitative analysis of this phenomenon. This mechanism can also explain the phenomena of slacking and force drop after turning visual feedback off during accurate force production task.