Phase transitions and postural deviations during bimanual kinesthetic tracking

Upper limb coordination was studied by examining pattern stability of between-hand rhythmical coordination. In the first of two experiments, relative phase of rhythmical wrist flexion-extension was examined within a kinesthetic tracking paradigm. Eight right-handed subjects actively tracked a driven hand being flexed and extended by a computer-controlled AC servo-motor. Hand movements were constrained in flexion or extension. The simultaneous contraction of wrist flexors and extensors was defined as inphase (IP) and the alternating contraction of wrist flexors and extensors as antiphase (AP). Phase transitions (from AP to IP) were observed in 16% of trials prepared in AP. Fewer phase transitions occurred when the right wrist was constrained in flexion, and also when the left wrist was constrained in extension. IP patterns were performed with greater stability than AP patterns. These effects were explored further in a second experiment with the addition of a secondary probe reaction time task to assess demands on central capacity, and the analysis of wrist flexor and extensor electromyographic activity. Subjects returned longer reaction times for AP than IP movement, suggesting the AP movement pattern placed a greater demand on central capacity than the IP movement pattern. During this kinesthetic tracking task, similar dynamic principles emerged as those observed during bilaterally active bimanual rhythmical coordination. The greater stability of the hand-posture combination where the driven left hand was constrained in extension and the active right hand was constrained in flexion may be a demonstration of unique central control of coupled activity. Electronic Publication     

Role of peripheral afference during acquisition of a complex coordination task

It has long been supposed that the interference observed in certain patterns of coordination is mediated, at least in part, by peripheral afference from the moving limbs. We manipulated the level of afferent input, arising from movement of the opposite limb, during the acquisition of a complex coordination task. Participants learned to generate flexion and extension movements of the right wrist, of 75 degrees amplitude, that were a quarter cycle out of phase with a 1-Hz sinusoidal visual reference signal. On separate trials, the left wrist either was at rest, or was moved passively by a torque motor through 50 degrees, 75 degrees or 100 degrees, in synchrony with the reference signal. Five acquisition sessions were conducted on successive days. A retention session was conducted 1 week later. Performance was initially superior when the opposite limb was moved passively than when it was static. The amplitude and frequency of active movement were lower in the static condition than in the driven conditions and the variation in the relative phase relation across trials was greater than in the driven conditions. In addition, the variability of amplitude, frequency and the relative phase relation during each trial was greater when the opposite limb was static than when driven. Similar effects were expressed in electromyograms. The most marked and consistent differences in the accuracy and consistency of performance (defined in terms of relative phase) were between the static condition and the condition in which the left wrist was moved through 50 degrees. These outcomes were exhibited most prominently during initial exposure to the task. Increases in task performance during the acquisition period, as assessed by a number of kinematic variables, were generally well described by power functions. In addition, the recruitment of extensor carpi radialis (ECR), and the degree of co-contraction of flexor carpi radialis and ECR, decreased during acquisition. Our results indicate that, in an appropriate task context, afferent feedback from the opposite limb, even when out of phase with the focal movement, may have a positive influence upon the stability of coordination.     

Reprogramming of muscle activation patterns at the wrist in compensation for elbow reaction torques during planar two-joint arm movements

The relationship between wrist kinematics, dynamics and the pattern of muscle activation were examined during a two-joint planar movement in which the two joints moved in opposite directions, i.e. elbow flexion/wrist extension and elbow extension/wrist flexion. Elbow movements (ranging from 10 to 70 deg) and wrist movements (ranging from 10 to 50 deg) were performed during a visual, step-tracking task in which subjects were required to attend to the initial and final angles at each joint. As the elbow amplitude increased, wrist movement duration increased and the wrist movement trajectories became quite variable. Analysis of the torques acting at the wrist joint showed that elbow movements produced reaction torques acting in the same direction as the intended wrist movement. Distinct patterns of muscle activation were observed at the wrist joint that were dependent on the relative magnitude of the elbow reaction torque in relation to the net wrist torque. When the magnitude of the elbow reaction torque was quite small, the wrist agonist was activated first. As the magnitude of the elbow reaction torque increased, activity in the wrist agonist decreased significantly. In conditions where the elbow reaction torque was much larger than the net wrist torque, the wrist muscle torque reversed direction to oppose the intended movement. This reversal of wrist muscle torque was directly associated with a change in the pattern of muscle activation where the wrist antagonist was activated prior to the wrist agonist. Our findings indicate that motion of the elbow joint is an important consideration in planning wrist movement. Specifically, the selection of muscle activation patterns at the wrist is dependent on the relative magnitude and direction of the elbow reaction torque in relation to the direction of wrist motion.     

Motor unit recruitment in human medial gastrocnemius muscle during combined knee flexion and plantarflexion isometric contractions

Previous work on multifunctional muscle has suggested that motor unit recruitment during a combined force task is the result of an interactive effect of weighted inputs acting simultaneously on the motoneuron pool. The present study shows that a similar effect describes motor unit activation in a two-joint muscle as forces are combined at both proximal and distal attachments. The recruitment thresholds of single motor units in medial gastrocnemius muscle were determined during combined knee flexion and plantarflexion isometric contractions. Slow isometric ramp contractions in knee flexion were produced while maintaining various background levels of plantarflexion force. The combination of knee flexion and plantarflexion forces at which a motor unit initially discharged was used to characterize recruitment as represented by the slope of the regression line fit to the individual data points. Each subject completed two experiments; one at each of two knee joint angles, with the ankle joint fixed at 90°. The effect of knee angle was assessed by comparing the slopes of the regression lines that characterized motor unit recruitment at each knee angle. Motor units in medial gastrocnemius were recruited when the linear sum of the forces exerted in plantarflexion and knee flexion exceeded a certain threshold of combined force. Specifically, the apparent force threshold of recruitment in knee flexion decreased as the level of force maintained in plantarflexion increased. Further, evidence is provided indicating that the linear relationship describing recruitment in two-joint muscle is dependent upon joint angle. The basis for the alteration in force threshold is thought to be related to changes in muscle length and mechanical advantage which might adjust the relative weighting of inputs that determine muscle activation patterns. These results indicate a possible common strategy employed by the nervous system in coordinating the activation of motor units to perform a specific task.     

A comprehensive analysis of muscle recruitment patterns during shoulder flexion: an electromyographic study

Although flexion is a common component of the routine clinical assessment of the shoulder the muscle recruitment patterns during this movement are not clearly understood making valid interpretation of potential muscle dysfunction problematic. The purpose of this study was to comprehensively examine shoulder muscle activity during flexion in order to compare the activity levels and recruitment patterns of shoulder flexor, scapular lateral rotator and rotator cuff muscles. Electromyographic (EMG) data were recorded from 12 shoulder muscles sites in 15 volunteers. Flexion was performed in standing in the sagittal plane at no load, 20%, and 60% of each subject's maximum load. EMG data were normalized to maximum values obtained during maximum voluntary contractions. Results indicated that anterior deltoid, pectoralis major, supraspinatus, infraspinatus, serratus anterior, upper, and lower trapezius were activated at similar moderate levels. However, subscapularis was activated at low levels and significantly lower than supraspinatus and infraspinatus. Similar activity patterns across time were demonstrated in the muscles that produce flexion torque, laterally rotate the scapula, as well as supraspinatus and infraspinatus, and did not change as flexion load increased. The onset of activity in supraspinatus and anterior deltoid occurred at the same time and prior to movement of the limb at all loads with infraspinatus activity also occurring prior to movement onset at the medium and high load conditions only. Posterior rotator cuff muscles appear to be counterbalancing anterior translational forces produced during flexion and it would appear that supraspinatus is one of the muscles that consistently "initiates" flexion.     

Forces applied by the incisors and roles of periodontal afferents during food-holding and -biting tasks

The force exerted by the central incisors while holding and splitting a food morsel was analyzed to characterize human biting behavior. The force was continuously sampled by a transducer-equipped plate upon which a small piece of dry biscuit or half a peanut rested. Subjects were instructed to position the plate between the incisor teeth and to split the morsel either immediately (split task) or after holding it for a brief period (hold-and-split task). While holding either food substance between the incisors, subjects automatically exerted light contact forces of less than 1 N (0.36–0.76N range among subjects). Considering that the subjects had no instructions about what force levels to employ, the hold force was remarkably stable during individual trials and highly similar among trials. Even during the split task, subjects opted to hold the morsel momentarily on ca. 50% of the trials with a similar, low contact force. For both tasks, subjects split the morsel by exerting a distinct, rapidly executed ramp increase in force. The split occurred at 7.8–10.3 N (range among subjects) bite force for the biscuit and 16.0–19.0 N for the peanut. The magnitude of the forces used during the hold phase were within the range over which most periodontal afferents are optimally sensitive to changes in force, i.e., forces below about 1 N. This observation suggested that the subjects automatically adjusted the force to maximize the availability of information from periodontal afferents and avoided higher forces at which the sensitivity of most afferents was not optimal. We further confirmed that the periodontal receptors serve a role in controlling the hold force by anesthetizing the periodontal tissues: subjects employed considerably higher and more variable hold forces, but there was no effect on the split phase. In addition, the morsel frequently escaped from the incisal edges of the teeth while the subject attempted to maintain it in position. It was concluded that subjects rely on signals from periodontal afferents to regulate the jaw muscles, particularly when they first contact, manipulate, and hold food substances between the teeth.     

Changes in muscle afferents, motoneurons and motor drive during muscle fatigue

Fatigue is a reduction of maximal muscle force or power that occurs with exercise. It is accompanied by changes at multiple levels in the motor pathway and also by changes in the discharge patterns of muscle afferents. Changes in afferent firing can lead to altered perceptions and can also act on the efferent pathway. Changes in the motor pathway include slowing of motor unit firing rates during sustained maximal voluntary contractions (MVCs). Muscle responses to stimulation at different levels of the motor pathway also change. Transcranial magnetic stimulation of the motor cortex and stimulation of descending tracts in the spinal cord in human subjects show an increase in the response of the cortex and a decrease in response of the motoneuron pool during sustained MVCs. In addition, the silent period following magnetic stimulation is prolonged. During relaxation after fatiguing exercise, muscle responses to stimulation of the motor cortex are initially facilitated and are then depressed for many minutes, whereas responses to descending tract stimulation are initially depressed but recover over about 2 min. Although some of the loss of force of fatigue does occur through inadequate drive to the muscle, it is not clear which, if any, of the changes described in the cortex or the motoneurons are responsible for loss of maximal voluntary force and thus contribute to fatigue. Changes may be associated with muscle fatigue without causing it. Accepted: 30 May 2000     

The effect of muscle length on motor-unit recruitment during isometric plantar flexion in humans

The triceps surae muscle group, consisting of the mono-articular soleus (SOL) and bi-articular gastrocnemius (GAS) muscles, primarily generates plantar flexor torque. Since the GAS muscle crosses the knee joint, flexion of the knee reduces the length of this muscle, thus limiting its contribution to torque output. However, it is not clearly understood how the central nervous system activates muscles that are at inefficient or non-optimal force-producing lengths. Therefore, the present study was designed to determine the effect of muscle length on motor-unit recruitment in the medial GAS muscle. Single motor-unit activity was recorded from the medial GAS muscle while electromyographic (EMG) activity was recorded from the SOL muscle in nine male subjects. With the ankle angle held constant at 90 degrees, the knee angle was changed from 180 degrees to 90 degrees, corresponding to a long and short GAS muscle length, respectively. Levels of voluntary plantar flexor torque were produced at a rate of 2 Nm.s-1 until motor-unit activity was detected. A total of 229 motor units were recorded, of which 121 and 108 were obtained at the long and short muscle lengths, respectively. At the short length, onset of motor-unit activity occurred at significantly higher levels of plantar flexor torque and SOL EMG. Onset of motor-unit activity occurred at 2.97 +/- 7.78 Nm and 32.14 +/- 10.25 Nm, corresponding to 0.045 +/- 0.075 mV and 0.231 +/- 0.129 mV of SOL EMG in the long and short positions, respectively. No individual GAS motor unit could be recorded at both muscle lengths. Motor units in the shortened GAS muscle may be influenced by peripheral afferents capable of reducing the excitability of the motoneurone pool. This may also reflect a specific inhibition of motor units having shortened, non-optimal fascicle lengths, and they are thereby incapable of contributing to plantar flexor torque.     

Changes in finger coordination and responses to single pulse TMS of motor cortex during practice of a multifinger force production task

We investigated the changes in finger coordination and in finger force responses to transcranial magnetic stimulation (TMS) applied over the motor cortex associated with a single practice session of an accurate ramp force production task. Subjects pressed with their index, middle and ring fingers onto three force transducers fixed to a rigid platform that was balanced on a narrow pivot under the middle finger. The task was to produce a smoothly increasing ramp of total force from 0 to 25 N over 4 s following a visual target. Subjects performed three brief series of trials without TMS (12 trials each) in the beginning, in the middle, and in the end of the experiment. The main part of the experiment involved 173 trials, and in each of them at random times in the ramp a suprathreshold TMS pulse was applied over the hand area of the contralateral motor cortex in order to evoke a twitch in the finger flexor muscles. At the end of the experiment the subjects also performed 12 constant force production trials, and TMS was unexpectedly applied in each trial. During the ramp force trials the amplitude of the response to TMS was largely independent of the force exerted at the time of stimulation, whereas in static holding trials the amplitude of the response increased with higher levels of background contraction. Over time subjects improved their overall tracking performance: the variance of the force trajectory (VarF_TOT), as computed over sets of unperturbed trials, declined by 60% after the first 100 trials, but there was little additional improvement after the second 100 trials. Variance in the force finger space related to the total moment with respect to the pivot also showed a decline during the first half of practice and minimal further changes during the second half. In contrast, finger force variance that did not affect either total force or total moment showed no changes after the first 100 trials and a decline during the second 100 trials. This variance component quantified per finger was significantly larger than those related to the total force and total moment. The mean size of the TMS-induced phasic force increment decreased by 12% over the course of the 200 trials. The forces evoked in the index and ring fingers gradually became more equal, reducing the total moment with respect to the pivot and improving balance. We speculate that development of a relatively low twitch force with low total moment on the pivot made it easier for subjects to continue tracking after the TMS pulse. Such changes could well be correlated with the degree of corticospinal involvement in the task. The results suggest task specific, practice-related plastic changes in neural structures involved in the responses to TMS.     

Lip muscle reflex and intentional response levels in a simple speech task

Summary Sensorimotor integration in human lip muscle was studied by recording muscle activity while subjects produced simple speech utterances in response to mechanical stimulation. On each trial subjects were instructed either to produce the syllable pa or not respond when they detected movement of a small paddle held between the lips. Mechanical stimuli were adequate to elicit reflexes over poststimulus intervals of 15–30 ms (R1) and 30–50 ms (R2). EMG recordings were obtained from upper and lower lip muscles, and EMG levels were calculated for individual trials over several poststimulus time intervals. The independent effects of stimulus magnitude, prestimulus EMG, and reaction time on poststimulus response levels were assessed using multiple regression analysis. R1 and R2 levels were positively correlated with stimulus magnitude, but stimulus magnitude had little modulating effect on intentional lip muscle responses. Both reflex and intentional response levels showed positive associations with prestimulus EMG level. Instructional set had significant modulating effects on reflex responses in 9 of 10 subjects, but the nature of these effects varied among subjects. These various findings are discussed in relation to similar studies on limb motor systems and lip motor control for speech.     

Coordinated isometric muscle commands adequately and erroneously programmed for the weight during lifting task with precision grip

Summary Small objects were lifted from a table, held in the air, and replaced using the precision grip between the index finger and thumb. The adaptation of motor commands to variations in the object's weight and sensori-motor mechanisms responsible for optimum performance of the transition between the various phases of the task were examined. The lifting movement involved mainly a flexion of the elbow joint. The grip force, the load force (vertical lifting force) and the vertical position were measured. Electromyographic activity (e.m.g.) was recorded from four antagonist pairs of hand/arm muscles primarily influencing the grip force or the load force. In the lifting series with constant weight, the force development was adequately programmed for the current weight during the loading phase (i.e. the phase of parallel increase in the load and grip forces during isometric conditions before the lift-off). The grip and load force rate trajectories were mainly single-peaked, bell-shaped and roughly proportional to the final force. In the lifting series with unexpected weight changes between lifts, it was established that these force rate profiles were programmed on the basis of the previous weight. Consequently, with lifts programmed for a lighter weight the object did not move at the end of the continuous force increase. Then the forces increased in a discontinous fashion until the force of gravity was overcome. With lifts programmed for a heavier weight, the high load and grip force rates at the moment the load force overcame the force of gravity caused a pronounced positional overshoot and a high grip force peak, respectively. In these conditions the erroneous programmed commands were automatically terminated by somatosensory signals elicited by the start of the movement. A similar triggering by somatosensory information applied to the release of programmed motor commands accounting for the unloading phase (i.e. the parallel decrease in the grip and load forces after the object contacted the table following its replacement). These commands were always adequately programmed for the weight.     

Bilateral deficit and symmetry in finger force production during two-hand multifinger tasks

A comprehensive study of patterns of finger forces during one-hand and two-hand multifinger maximal force production trials was performed with particular emphasis on differences between tasks involving symmetrical and asymmetrical finger groups (symmetrical and asymmetrical tasks). Twelve healthy right-handed subjects performed maximal voluntary force production tasks with different finger combinations. Force deficit (FD) for a finger group within a hand was defined as a drop in peak force in a multifinger task as compared to the sum of individual finger peak forces in single-finger tasks. FD showed a dependence on both the number of fingers within the hand and the number of fingers in the other hand. An additional drop in peak finger forces was seen in two-hand tests (bilateral deficit, BD). BD summed over two hands was independent of the number of fingers involved in the two-hand tasks, but dependent on the distribution of fingers between the two hands. BD for a hand was larger for tasks involving fewer fingers within the hand and more fingers in the other hand. It was higher for asymmetrical tasks than for symmetrical tasks. The difference between asymmetrical and symmetrical tasks was due to the different behavior of asymmetrically involved fingers. FD was larger for asymmetrical master (explicitly involved) fingers, while forces produced involuntarily by asymmetrical slave (explicitly non-involved) fingers were larger. These differences brought down the total moment produced by both hands in the frontal plane. FD and BD are phenomena of different origin whose effects sum up. The observations have led to further development of a previously proposed double-representation, mirror-image (DoReMi) hypothesis and refinement of the neural network underlying the two-hand finger interaction.     

GABAergic and glycinergic neural inhibition in excitatory frequency tuning of bat inferior collicular neurons

This study examines the impact of peripheral nerve block, that is, the elimination of tactile feedback on synchronization performance. In a tapping experiment in which subjects were instructed to tap in synchrony with an auditory pacing signal, three different tasks were studied under conditions with and without peripheral nerve block: standard tapping with tactile contact, isometric tapping, and contact-free tapping. In addition, the maximum tapping rate was registered both with and without peripheral nerve block. It was found that the anticipatory error, usually observed in synchronization tasks, was affected by the peripheral nerve block in the standard tapping and the isometric tapping task. In both tasks, local anesthesia led to an increase in asynchrony between the pacing signal and the tap. Performance remained unimpaired in those tasks in which tactile information was assumed to play a minor role (maximum tapping rate and contact-free tapping). The results clearly demonstrate the importance of tactile feedback for the timing of movements. The predictions of a model assuming a strong correlation between the amount of sensory feedback and the size of the negative asynchrony in synchronization tasks were examined and discussed.     

Control of grip force during restraint of an object held between finger and thumb: responses of muscle and joint afferents from the digits

Pulling or pushing forces applied to an object gripped between finger and thumb excite tactile afferents in the digits in a manner awarding these afferents probable roles in triggering the reactive increases in grip force and in scaling the changes in grip force to the changes in applied load-force. In the present study we assessed the possible contributions from slowly adapting afferents supplying muscles involved in the generation of grip forces and from digital joint afferents. Impulses were recorded from single afferents via tungsten microelectrodes inserted percutaneously into the median or ulnar nerves of awake human subjects. The subject held a manipulandum with a precision grip between the receptor-related digit (index finger, middle finger, ring finger or thumb) and an opposing digit (thumb or index finger). Ramp-and-hold load forces of various amplitudes (0.5–2.0 N) and ramp rates (2–32 N/s) were delivered tangential to the parallel grip surfaces in both the distal (pulling) and the proximal (pushing) directions. Afferents from the long flexors of the digits (n=19), regardless of their muscle-spindle or tendon-organ origin, did not respond to the load forces before the onset of the automatic grip response, even with the fastest ramp rates. Their peak discharge closely followed the peak rate of increase in grip force. During the hold phase of the load stimulus, the afferents sustained a tonic discharge. The discharge rates were significantly lower with proximally directed loads despite the mean grip-force being similar in the two directions. This disparity could be explained by the differing contributions of these muscles to the finger-tip forces necessary to restrain the manipulandum in the two directions. Most afferents from the short flexors of the digits (n=17), including the lumbricals, dorsal interossei, opponens pollicis, and flexor pollicis brevis, did not respond at all, even with the fastest ramps. Furthermore, the ensemble pattern from the joint afferents (n=6) revealed no significant encoding of changes in finger-tip forces before the onset of the increase in grip force. We conclude that mechanoreceptors in the flexors of the digits and in the interphalangeal joints cannot be awarded a significant role in triggering the automatic changes in grip force. Rather, their responses appeared to reflect the reactive forces generated by the muscles to restrain the object. Hence, it appears that tactile afferents of the skin in contact with the object are the only species of receptor in the hand capable of triggering and initially scaling an appropriate change in grip force in response to an imposed change in load force, but that muscle and joint afferents may provide information related to the reactive forces produced by the subject.     

The role of predictive visual temporal information in the coordination of muscle activity in catching

Summary This study addresses the question as to the nature of the information on which the preactivation of the appropriate muscles in the grasping of the ball in a onehanded catching task is initiated and coordinated. High speed film and electromyograms were recorded while experiences subjects (N = 4) caught balls — projected towards them by a ball-machine at different speeds (11.9, 13.9 and 16.2 m/s — resulting in significantly different flight times of 508, 443 and 355 ms, respectively). Tau-margins (times to contact) values were calculated at the time of the initiation of the grasp movement for each subject at each speed. No significant differences were found between taumargins at different speeds. Further, the onset of the muscle activity for the initiation of the grasp movement was shown to be independent of ball speed. These findings lend support to the contention that the initiation of the grasp movement in catching is controlled and coordinated by the optical variable tau which specifies (directly) this time-to-contact. Given that the muscle group selected includes both flexors and extensors, co-activation on the basis of tau information is evidenced.     

Changes in the functional MR signal in motor and non-motor areas during intermittent fatiguing hand exercise

The aim of this study was to determine whether there were significant changes in the time course of the functional magnetic resonance imaging (fMRI) signal in motor and non-motor regions of both cerebral hemispheres during a unilateral fatiguing exercise of the hand. Twelve subjects performed a submaximal (30%) intermittent fatiguing handgrip exercise (3 s grip, 2 s release, left hand) for ∼9 min during fMRI scanning. Regression analysis was used to measure changes in fMRI signal from primary sensorimotor cortex (SM1), premotor cortex and visual cortex (V1) in both hemispheres. Force declined to 77 ± 3.6% of prefatigue maximal force (P < 0.05). The fMRI signal from SM1 contralateral to the fatiguing hand increased by 1.2 ± 0.5% of baseline (P < 0.05). The fMRI signal from the ipsilateral SM1 did not change significantly. Premotor cortex showed a similar pattern but did not reach significance. The signal from V1 increased significantly for both hemispheres (contralateral 1.3 ± 0.9%, ipsilateral 1.5 ± 0.9% of baseline and P < 0.05). During the performance of a unimanual, submaximal fatiguing exercise there is an increase in activation of motor and non-motor regions. The results are in keeping with the notion of an increase in sensory processing and corticomotor drive during fatiguing exercise to maintain task performance as fatigue develops.     

Coherence analysis of muscle activity during quiet stance

Studies of muscle activation during perturbed standing have demonstrated that the typical patterns of coordination (“ankle strategy” and “hip strategy”) are controlled through multiple muscles activated in a distal-to-proximal or proximal-to-distal temporal pattern. In contrast, quiet stance is thought to be maintained primarily through the ankle musculature. Recently, spectral analysis of inter-segment body motion revealed the coexistence of both ankle and hip patterns of coordination during quiet stance, with the predominating pattern dependent on the frequency of body sway. Here we use frequency domain techniques to determine if these patterns are associated with the same muscular patterns as observed during perturbed stance. Six of the seven muscles measured showed a linear relationship to the sway of at least one body segment, all being leg muscles. Muscle–segment phases were consistent with that required to resist gravity at low frequencies, with increasing phase lag as frequency increased. Visual information had effects only at frequencies below 0.5 Hz, where the shift from in-phase to anti-phase trunk–leg co-phase was observed. These results indicate that co-existence of the ankle and hip pattern during quiet stance involves only leg musculature. Anti-phase movement of the trunk relative to the legs at higher frequencies arises from indirect biomechanical control from posterior leg muscles.     

Force adaptation transfers to untrained workspace regions in children

When humans perform goal-directed arm movements under the influence of an external damping force, they learn to adapt to these external dynamics. After removal of the external force field, they reveal kinematic aftereffects that are indicative of a neural controller that still compensates the no longer existing force. Such behavior suggests that the adult human nervous system uses a neural representation of inverse arm dynamics to control upper-extremity motion. Central to the notion of an inverse dynamic model (IDM) is that learning generalizes. Consequently, aftereffects should be observable even in untrained workspace regions. Adults have shown such behavior, but the ontogenetic development of this process remains unclear. This study examines the adaptive behavior of children and investigates whether learning a force field in one hemifield of the right arm workspace has an effect on force adaptation in the other hemifield. Thirty children (aged 6-10 years) and ten adults performed 30 degrees elbow flexion movements under two conditions of external damping (negative and null). We found that learning to compensate an external damping force transferred to the opposite hemifield, which indicates that a model of the limb dynamics rather than an association of visited space and experienced force was acquired. Aftereffects were more pronounced in the younger children and readaptation to a null-force condition was prolonged. This finding is consistent with the view that IDMs in children are imprecise neural representations of the actual arm dynamics. It indicates that the acquisition of IDMs is a developmental achievement and that the human motor system is inherently flexible enough to adapt to any novel force within the limits of the organism's biomechanics.     

Changes in rolandic mu rhythm during observation of a precision grip

We recorded 128-channel EEG from 16 participants while they observed, imitated, and self-initiated the precision grip of a manipulandum. Mu rhythm amplitudes were significantly lower during observation of a precision grip than during observation of a simple hand extension without object interaction. Scalp topographies for subtractions of observation, imitation, and execution conditions from the control condition showed a high degree of congruence, supporting the notion of a human observation-execution matching system. Surface Laplacian transformations suggest that the decrease in mu amplitude during precision grip observation reflects desynchronization of mu rhythm generators in the sensorimotor cortex. These results support the hypothesis that sensorimotor cortex is a neural substrate involved in the representation of both self- and other-generated actions and show the mu rhythm is sensitive to subtle changes in observed motor behavior.     

Effects of varying task constraints on solutions to joint coordination in a sit-to-stand task

The question of how multijoint movement is controlled can be studied by discovering how the variance of joint trajectories is structured in relation to important task-related variables. In a previous study of the sit-to-stand task, for instance, variations of body segment postures that leave the position of the body's center of mass (CM) unchanged were significantly greater than variations of body segment posture that varied the CM position. The present experiments tested the hypothesis that such structuring of joint configuration variability is accentuated when the mechanical or perceptual task demands are made more challenging. Six subjects performed the sit-to-stand task without vision (eyes closed), either on a normal or on a narrow support surface. An additional constraint on the postural task was introduced in a third condition by requiring subjects to maintain light touch (less than 1 N) with the fingertips while coming to a standing position on the narrow base of support. The joint configurations observed at each point in normalized time were analyzed with respect to trial-to-trial variability. The task variables CM and head position were used to define goal-equivalent sets of joint configurations ("uncontrolled manifolds," UCMs) within which variation of joint configuration leaves the task variables unchanged. The variability of joint configurations across trials was decomposed into components that did not affect (within the UCM) and that did affect (orthogonal to the UCM) the values of these task variables. Our results replicate the earlier finding of much larger variability in directions of joint space that leave the CM unchanged compared with directions that affect CM position. This effect was even more pronounced here than in the previous experiment, probably because of the more difficult perceptual conditions in the current study (eyes closed). When the mechanical difficulty of the task was increased, the difference between the two types of joint variability was further accentuated, primarily through increase in goal-equivalent variance. This provides evidence for the hypothesis that under challenging task constraints increased variability is selectively directed into task-irrelevant degrees of freedom. Because differential control along different directions of joint space requires coordination among joint angles, this observation supports the view that the CNS responds to increased task difficulty through enhanced coordination among degrees of freedom. The adaptive nature of this coordination is further illustrated by the similar enhanced use of goal-equivalent joint combinations to achieve a stable CM position when subjects stood up under the additional constraint of maintaining light touch with the fingertips. This was achieved by channeling goal-equivalent variability into different directions of joint configuration space.