Event-related beta EEG changes during wrist movements induced by functional electrical stimulation of forearm muscles in man

Event-related beta electroencephalographic (EEG) changes were studied during wrist movements induced by functional electrical stimulation (FES) of the appropriate forearm muscles in healthy volunteers. Active and passive hand movements were investigated as control conditions. Significant EEG changes with respect to a pre-movement period were analyzed by calculating time-frequency maps of event-related (de-)synchronization (ERD/ERS) for 32 EEG channels recorded from sensorimotor and premotor areas. Immediately after the beginning of the FES movement, a prominent ERD was found, followed by a beta ERS similar to that observed after active or passive wrist movements. Both changes were maximal over the contralateral primary hand area. The main difference between active and stimulation-induced movements was that in the latter case no ERD was detectable prior to movement-onset. These findings suggest that the sensorimotor processing during FES involves some of the processes which are also involved in voluntary hand movements.     

Human motor compensations for thixotropy-dependent changes in muscular resting tension after moderate joint movements

AIM: This study on healthy subjects explores history-dependent changes in the resting tension of relaxed wrist muscles after moderate joint excursions and the motor control consequences of these changes during voluntary wrist joint position maintenance. METHODS: Integrated surface electromyogram (IEMG) was recorded from wrist extensor/flexor muscles. Angular position and torque were recorded from the wrist joint. Changes in wrist flexor muscle resting tension were sensed by a force transducer pressed against the tendons. RESULTS: Consecutive stepwise changes (7.5 degrees ) in wrist joint position (within the dorsiflexed range) were either imposed on relaxed subjects or actively performed while the subjects under visual guidance tried to mimic the passive movements. In relaxed subjects, passive joint torque resistance at a given steady dorsiflexed position either gradually declined or rose depending on the direction of the previous transition movements. In corresponding voluntary contraction experiments, the IEMG amplitude from position holding wrist extensors was found to vary in a similar way as the passive torque resistance. Further, there was a strong correlation between history-dependent changes in extensor IEMG amplitude and stress alterations exhibited by the relaxed antagonist flexors. The above described, slowly subsiding post-movement mechanical and motor adaptations were accelerated by brief forceful cocontractions of the forearm muscles. CONCLUSION: Moderate stepwise changes in joint position are sufficient to induce history-dependent after-effects in passive muscular resting tension, after-effects which during voluntary position holding are effectively compensated for by the motor control system.     

Human motor compensations for thixotropy‐dependent changes in muscular resting tension after moderate joint movements

Aim: This study on healthy subjects explores history‐dependent changes in the resting tension of relaxed wrist muscles after moderate joint excursions and the motor control consequences of these changes during voluntary wrist joint position maintenance. Methods: Integrated surface electromyogram (IEMG) was recorded from wrist extensor/flexor muscles. Angular position and torque were recorded from the wrist joint. Changes in wrist flexor muscle resting tension were sensed by a force transducer pressed against the tendons. Results: Consecutive stepwise changes (7.5°) in wrist joint position (within the dorsiflexed range) were either imposed on relaxed subjects or actively performed while the subjects under visual guidance tried to mimic the passive movements. In relaxed subjects, passive joint torque resistance at a given steady dorsiflexed position either gradually declined or rose depending on the direction of the previous transition movements. In corresponding voluntary contraction experiments, the IEMG amplitude from position holding wrist extensors was found to vary in a similar way as the passive torque resistance. Further, there was a strong correlation between history‐dependent changes in extensor IEMG amplitude and stress alterations exhibited by the relaxed antagonist flexors. The above described, slowly subsiding post‐movement mechanical and motor adaptations were accelerated by brief forceful cocontractions of the forearm muscles. Conclusion: Moderate stepwise changes in joint position are sufficient to induce history‐dependent after‐effects in passive muscular resting tension, after‐effects which during voluntary position holding are effectively compensated for by the motor control system.     

Properties of human spinal interneurones: normal and dystonic control

The muscles that control wrist posture receive large inputs from reflexes driven by hand afferents. In several studies, we have investigated these reflexes by electrical stimulation of cutaneous (median nerve) and proprioceptive (ulnar nerve) afferents from the hand. Median stimulation produced short latency inhibition in all motor nuclei investigated, possibly through inhibitory propriospinal-like interneurones. Ulnar stimulation produced similar inhibition but only in wrist extensors. In the other motor nuclei, ulnar stimulation produced short latency excitation mediated by group I motoneuronal drive through both monosynaptic and non-monosynaptic pathways involving excitatory propriospinal-like interneurones. This was followed by late excitations mediated through spinal group II and trans-cortical group I pathways. These results show that these pathways are concerned with the integration of afferent inputs, proprioceptive and cutaneous, to control of wrist posture during hand movements. Patients with focal hand dystonia exhibit abnormal postures. To investigate whether these spinal pathways contribute to these conditions, the effects of ulnar stimulation on wrist muscle activity during voluntary tonic contraction were examined in patients who suffer writer's cramp. Ulnar-induced inhibition of the wrist extensors was reduced on the dystonic side of patients compared with their normal side and controls. In patients who exhibited abnormal wrist posture, group II excitation of the wrist flexors was also modified on the dystonic side. Cutaneous stimuli, by contrast, increased wrist flexor EMG on both sides and only in patients who exhibited normal posture. We conclude that spinal interneurones have a significant role in integrating afferent inputs from the hand to control wrist posture during hand movements and that altered function in these spinal networks is associated with the complex pathophysiology of writer's cramp.     

Human finger independence: limitations due to passive mechanical coupling versus active neuromuscular control

We studied the extent to which mechanical coupling and neuromuscular control limit finger independence by studying passive and active individuated finger movements in healthy adults. For passive movements, subjects relaxed while each finger was rotated into flexion and extension by a custom-built device. For active movements, subjects moved each finger into flexion and extension while attempting to keep the other, noninstructed fingers still. Active movements were performed through approximately the same joint excursions and at approximately the same speeds as the passive movements. We quantified how mechanical coupling limited finger independence from the passive movements, and quantified how neuromuscular control limited finger independence using an analysis that subtracted the indices obtained in the passive condition from those obtained in the active condition. Finger independence was generally similar during passive and active movements, but showed a trend toward less independence in the middle, ring, and little fingers during active, large-arc movements. Mechanical coupling limited the independence of the index, middle, and ring fingers to the greatest degree, followed by the little finger, and placed only negligible limitations on the independence of the thumb. In contrast, neuromuscular control primarily limited the independence of the ring, and little fingers during large-arc movements, and had minimal effects on the other fingers, especially during small-arc movements. For the movement conditions tested here, mechanical coupling between the fingers appears to be a major factor limiting the complete independence of finger movement.     

Implant provision of key,pinch and power grips in a C6 tetraplegic

An 11-channel multiplexed stimulator of nerves and muscles in the left forearm was implanted for hand control in January 1986 in a 21 year old woman who, after sustaining a C6 spinal lesion 7 years earlier, had voluntary shoulder and elbow movement but paralysed hands, trunk and legs. The patient controls the stimulation via a microcomputer control box and an RF transdermal link. We have investigated the control of her stimulated hand with a joystick under her contralateral hand which she moves from the shoulder and elbow. Since 1986, we have tried a variety of joystick control schemes involving power and key grips. Currently, for grip adjustment, forward and backward joystick movements correspond to thumb extension and abduction, respectively giving in addition both finger and wrist extension, whereas right and left joystick movements yield first closure and thumb opposition/adduction and flexion, respectively. Useful grasps are available by moving the joystick forward and then left (key grip), by moving the joystick backward and left (pinch grip), or by moving the joystick back and right (power grip). Thus, three distinct grips may be selected using these three quadrants of joystick movement. An additional control mode was found to be desirable to augment the patient's limited voluntary wrist positioning and provide wrist stability while adjusting finger grip.     

Cerebellar nuclear cell activity during antagonist cocontraction and reciprocal inhibition of forearm muscles

Monkeys were trained to exert a maintained isometric pinch with the thumb and forefinger. This task reliably elicited a simultaneous cocontraction of the forearm muscles. The same monkeys were also taught to insert the open hand into a manipulandum, flex and extend the wrist 35 and 15 degrees, respectively, and maintain an isometric wrist position against a mechanical stop for 1 s. This second task comprised two conditions: a dynamic or movement phase and a static or isometric phase. Movement always involved a wrist displacement of 50 degrees. Although some forearm muscles demonstrated bidirectional activity during the wrist displacement phase, all the wrist and finger muscles were alternatively active in isometric flexion or extension. Of the neurons in the dentate and interposed nuclei that consistently changed discharge during repeated isometric prehension, over 90% (61/67) of the neurons increased activity during this cocontraction of forearm muscles. About 70% (47/67) of these same nuclear cells discharged with a reciprocal pattern of firing during alternating wrist flexion-extension movements. Forty-six neurons had sustained and reciprocal discharge during the maintained isometric wrist postures. No differences were seen between the activity patterns of dentate and interposed cells with respect to either the prehension task or the reciprocal wrist-movement task. The discharge frequency of some dentate and interpositus neurons could be correlated with prehensile force as well as velocity of wrist movement and torque developed by wrist muscles. Correlation coefficients were calculated between nuclear cell discharge and the amplitude of the surface EMGs of the flexors and extensors of the wrist and fingers during the wrist-movement task. Sixteen nuclear cells showed low-order, but reliably positive, correlations with one of the two forearm muscle groups (mean r = 0.33). In contrast, a sample of seven Purkinje cells recorded during the same task demonstrated low-order correlations that were negative in sign (mean r = -0.30) between discharge frequency and one of the two forearm EMGs.     

Coupled bilateral movements and active neuromuscular stimulation: intralimb transfer evidence during bimanual aiming

Motor improvements in chronic stroke recovery accrue from coupled protocols of bilateral movements and active neuromuscular stimulation. This experiment investigated coupled protocols and within-limb transfer between distal and proximal joint combinations. The leading question focused on within-limb transfer of coupled protocols on distal joints to a bimanual aiming task that involved proximal joints. Twenty-six volunteers completed one of three motor recovery protocols according to group assignments: (1) coupled bilateral involved concurrent wrist/finger movements on the unimpaired limb coupled with active stimulation on the impaired limb; (2) unilateral/active stimulation involved neuromuscular electromyogram-triggered stimulation on the impaired wrist/fingers; and (3) no protocol (control group). During the pretest and posttest, subjects performed transverse plane target aiming movements (29 cm) with vision available. The coupled bilateral group showed positive intralimb transfer post-treatment when both arms moved simultaneously. During the posttest, the coupled bilateral group displayed improved movement time, higher peak limb velocity, less variability in peak velocity, and less percentage of total movement time in the deceleration phase than during the pretest. The evidence confirms that within-limb transfer from distal joint training to proximal joint combinations is viable and generalizable in chronic stroke rehabilitation. Moreover, these intralimb transfer findings extend the evidence favoring motor improvements for coupled bilateral protocols during chronic stroke.     

Co-ordination of breathing with rhythmic head and eye movements and with passive turnings of the body

In the present study, we investigated co-ordination of breathing with active horizontal head and eye movements and with passive body turnings. The main purpose was to compare co-ordination with active voluntary and with passive movements. Co-ordination was defined according to the relative phases (RPs) between breathing and additional rhythms. It was present when at least five consecutive RPs scattered less than 25% of breath duration or at least three consecutive RPs scattered less than 10%. Additionally, we studied the influence of factors such as size of active muscle groups, mechanical interactions or involuntary motor activities. In consecutive experimental conditions, 17 healthy volunteers performed voluntary rhythmic head (with eyes open and closed) and eye movements and were moved passively (also with eyes open and closed). Co-ordination occurred in all experimental conditions. The highest percentage of co-ordinated breaths was found during active head movements with closed eyes but the difference with other conditions was not significant. However, periods of stable entrainment were observed only in this condition. The results demonstrate that co-ordination is possible even in the absence of voluntary rhythmic movements. Size of active muscle groups or mechanical interactions do not necessarily affect the degree of co-ordination. On the other hand, involuntary motor activities such as reflex eye movements associated with vestibular stimulation may have a considerable influence on co-ordination between other motor processes. These results strongly support the theory of co-ordination as an unconscious nervous interaction that is not based on mechanical or intentional factors.     

Command control for functional electrical stimulation hand grasp systems using miniature accelerometers and gyroscopes

Recent commercially available miniature sensors have the potential to improve the functions of functional electrical stimulation (FES) systems in terms of control, reliability and robustness. A new control approach using a miniature gyroscope and an accelerometer was studied. These sensors were used to detect the linear acceleration and angular velocity of residual voluntary movements on upper limbs and were small and easy to put on. Five healthy subjects and three cervical spinal cord injured subjects were recruited to evaluate this controller. Sensors were placed on four locations: the shoulder, upper arm, wrist and hand. A quick forward-and-backward movement was employed to produce a distinctive waveform that was different from general movements. A detection algorithm was developed to generate a command signal by identifying this distinctive waveform through the detection of peaks and valleys in the sensor's signals. This command signal was used to control different FES hand grasp patterns. With a specificity of 0.9, the sensors had a success rate of 85–100% on healthy subjects and 82–97% on spinal cord injured subjects. In terms of sensor placement, the gyroscope was better as a control source than the accelerometer for wrist and hand positions, but the reverse was true for the shoulder.     

Functional magnetic resonance adaptation reveals the involvement of the dorsomedial stream in hand orientation for grasping

Reach-to-grasp actions require coordination of different segments of the upper limbs. Previous studies have examined the neural substrates of arm transport and hand grip components of such actions; however, a third component has been largely neglected: the orientation of the wrist and hand appropriately for the object. Here we used functional magnetic resonance imaging adaptation (fMRA) to investigate human brain areas involved in processing hand orientation during grasping movements. Participants used the dominant right hand to grasp a rod with the four fingers opposing the thumb or to reach and touch the rod with the knuckles without visual feedback. In a control condition, participants passively viewed the rod. Trials in a slow event-related design consisted of two sequential stimuli in which the rod orientation changed (requiring a change in wrist posture while grasping but not reaching or looking) or remained the same. We found reduced activation, that is, adaptation, in superior parieto-occipital cortex (SPOC) when the object was repeatedly grasped with the same orientation. In contrast, there was no adaptation when reaching or looking at an object in the same orientation, suggesting that hand orientation, rather than object orientation, was the critical factor. These results agree with recent neurophysiological research showing that a parieto-occipital area of macaque (V6A) is modulated by hand orientation during reach-to-grasp movements. We suggest that the human dorsomedial stream, like that in the macaque, plays a key role in processing hand orientation in reach-to-grasp movements.     

Selectivity of voluntary finger flexion during ischemic nerve block of the hand

During ischemic nerve block of an extremity the cortical representations of muscles proximal to the block are known to expand, increasing the overlap of different muscle representations. Such reorganization mimics that seen in actual amputees. We investigated whether such changes degrade voluntary control of muscles proximal to the block. Nine subjects produced brief, isometric flexion force selectively with each fingertip before, during, and after ischemic block at the wrist. We recorded the isometric force exerted at the distal phalanx of each digit, along with electromyographic (EMG) activity from intrinsic and extrinsic finger muscles. Despite paralysis of the intrinsic hand muscles, and associated decrements in the flexion forces exerted by the thumb, index, and little fingers, the selectivity of voluntary finger flexion forces and of EMG activity in the extrinsic finger muscles that generated these forces remained unchanged. Our observations indicate that during ischemic nerve block reorganization does not eliminate or degrade motor representations of the temporarily deafferented and paralyzed fingers.     

Wrist muscle activation patterns and stiffness associated with stable and unstable mechanical loads

Summary The objectives of this study were to examine the effects of load mechanical characteristics and agonist-antagonist muscle cocontraction, on the stretch reflex response of wrist flexor muscles, and to measure the associated wrist stiffness. Subjects were required to maintain a constant wrist angle while operating against flexor loads with different stability characteristics (constant, elastic or unstable). We measured the stretch reflex responses and joint stiffness by applying step displacements of 3° and 10°. Subjects used very little cocontraction of wrist flexor and extensor muscles when the load was constant or elastic, but increased cocontraction dramatically when the load was unstable, in order to increase the wrist stiffness. Although the magnitude of stretch reflex responses also increased with cocontraction, this simply reflected the level of tonic flexor muscle activity. We found no evidence to suggest that phasic stretch reflexes contributed significantly to the joint stiffness in this task. Clear differences in flexor muscle synergy were observed in the presence and absence of cocontraction, particularly when comparing the FCR and FCU muscles.     

The passive stiffness of the wrist and forearm

Because wrist rotation dynamics are dominated by stiffness (Charles SK, Hogan N. J Biomech 44: 614-621, 2011), understanding how humans plan and execute coordinated wrist rotations requires knowledge of the stiffness characteristics of the wrist joint. In the past, the passive stiffness of the wrist joint has been measured in 1 degree of freedom (DOF). Although these 1-DOF measurements inform us of the dynamics the neuromuscular system must overcome to rotate the wrist in pure flexion-extension (FE) or pure radial-ulnar deviation (RUD), the wrist rarely rotates in pure FE or RUD. Instead, understanding natural wrist rotations requires knowledge of wrist stiffness in combinations of FE and RUD. The purpose of this report is to present measurements of passive wrist stiffness throughout the space spanned by FE and RUD. Using a rehabilitation robot designed for the wrist and forearm, we measured the passive stiffness of the wrist joint in 10 subjects in FE, RUD, and combinations. For comparison, we measured the passive stiffness of the forearm (in pronation-supination), as well. Our measurements in pure FE and RUD agreed well with previous 1-DOF measurements. We have linearized the 2-DOF stiffness measurements and present them in the form of stiffness ellipses and as stiffness matrices useful for modeling wrist rotation dynamics. We found that passive wrist stiffness was anisotropic, with greater stiffness in RUD than in FE. We also found that passive wrist stiffness did not align with the anatomical axes of the wrist; the major and minor axes of the stiffness ellipse were rotated with respect to the FE and RUD axes by ～20°. The direction of least stiffness was between ulnar flexion and radial extension, a direction used in many natural movements (known as the "dart-thrower's motion"), suggesting that the nervous system may take advantage of the direction of least stiffness for common wrist rotations.     

Development of an MR-compatible hand exoskeleton that is capable of providing interactive robotic rehabilitation during fMRI imaging

Following advances in robotic rehabilitation, there have been many efforts to investigate the recovery process and effectiveness of robotic rehabilitation procedures through monitoring the activation status of the brain. This work presents the development of a two degree-of-freedom (DoF) magnetic resonance (MR)-compatible hand device that can perform robotic rehabilitation procedures inside an fMRI scanner. The device is capable of providing real-time monitoring of the joint angle, angular velocity, and joint force produced by the metacarpophalangeal (MCP) and proximal interphalangeal (PIP) joints of four fingers. For force measurement, a custom reflective optical force sensor was developed and characterized in terms of accuracy error, hysteresis, and repeatability in the MR environment. The proposed device consists of two non-magnetic ultrasonic motors to provide assistive and resistive forces to the MCP and PIP joints. With actuation and sensing capabilities, both non-voluntary–passive movements and active–voluntary movements can be implemented. The MR compatibility of the device was verified via the analysis of the signal-to-noise ratio (SNR) of MR images of phantoms. SNR drops of 0.25, 2.94, and 11.82% were observed when the device was present but not activated, when only the custom force sensor was activated, and when both the custom force sensor and actuators were activated, respectively.     

Decoding M1 neurons during multiple finger movements

We tested several techniques for decoding the activity of primary motor cortex (M1) neurons during movements of single fingers or pairs of fingers. We report that single finger movements can be decoded with >99% accuracy using as few as 30 neurons randomly selected from populations of task-related neurons recorded from the M1 hand representation. This number was reduced to 20 neurons or less when the neurons were not picked randomly but selected on the basis of their information content. We extended techniques for decoding single finger movements to the problem of decoding the simultaneous movement of two fingers. Movements of pairs of fingers were decoded with 90.9% accuracy from 100 neurons. The techniques we used to obtain these results can be applied, not only to movements of single fingers and pairs of fingers as reported here, but also to movements of arbitrary combinations of fingers. The remarkably small number of neurons needed to decode a relatively large repertoire of movements involving either one or two effectors is encouraging for the development of neural prosthetics that will control hand movements.     

Neurorehabilitation with new functional electrical stimulation for hemiparetic upper extremity in stroke patients.

In recent years, our understanding of motor learning, neuroplasticity, and functional recovery after the occurrence of brain lesion has grown significantly. New findings in basic neuroscience have stimulated research in motor rehabilitation. Repeated motor practice and motor activity in a real-world environment have been identified in several prospective studies as favorable for motor recovery in stroke patients. Electrical stimulation can be applied in a variety of ways to the hemiparetic upper extremity following stroke. In this paper, an overview of current research into clinical and therapeutic applications of functional electrical stimulation (FES) is presented. In particular, electromyography (EMG)-initiated electrical muscle stimulation--but not electrical muscle stimulation alone--improves the motor function of the hemiparetic arm and hand. Triggered electrical stimulation is reported to be more effective than untriggered electrical stimulation in facilitating upper extremity motor recovery following stroke. Power-assisted FES induces greater muscle contraction by electrical stimulation in proportion to the voluntary integrated EMG signal picked up, which is regulated by a closed-loop control system. Power-assisted FES and motor point block for antagonist muscles have been applied with good results as a new hybrid FES therapy in an outpatient rehabilitation clinic for patients with stroke. Furthermore, a daily home program therapy with power-assisted FES using new equipment has been able to effectively improve wrist and finger extension and shoulder flexion. Proprioceptive sensory feedback might play an important role in power-assisted FES therapy. Although many physiotherapeutic modalities have been established, conclusive proof of their benefit and physiological models of their effects on neuronal structures and processes are still missing. A multichannel near-infrared spectroscopy study to noninvasively and dynamically measure hemoglobin levels in the brain during functional activity has shown that cerebral blood flow in the sensory-motor cortex on the injured side is higher during a power-assisted FES session than during simple active movement or simple electrical stimulation. Nevertheless, evidence-based strategies for motor rehabilitation are more easily available, particularly for patients with hemiparesis.     

Attenuation of pathological tremors by functional electrical stimulation I: Method

In this study we explored the possibility of suppressing pathological tremors using closed-loop functional electrical stimulation (FES) to activate the tremorogenic muscles out-of-phase. A displacement signal monitored with a transducer was filtered so as to be “tuned” to the tremor frequency at the wrist or elbow. The filtered signal was used to amplitude-modulate the electrical stimulation. The design process was based on measurements of the open-loop frequency response characteristics of the forearm and hand to stimulation of the elbow and wrist flexors and extensors in a number of subjects. These data allowed us to identify closed-loop configurations, which attenuated 2–5 Hz tremors substantially, while only minimally attenuating functional movements in the 0–1 Hz range. There was a fairly delicate balance between efficacy and the risk of instability. However, designs were identified that offered enough tremor suppression and adequate immunity to muscle/load variations for the technique to be considered seriously for clinical application.     

Interjoint coordination during handwriting-like movements

The present study investigates intrinsic preferences and tendencies in coordination of the wrist and finger movements during handwriting-like tasks. Movement of the inkless pen tip in nine right-handed subjects was registered with a digitizer. One circle-drawing task and four line-drawing tasks were included in the experiment. The line-drawing task included: (1) drawing with the wrist only, (2) drawing with the fingers only, (3) an equivalent pattern consisting of the simultaneous flexion/extension of the wrist and fingers, and (4) a nonequivalent pattern in which wrist flexion was accompanied by finger extension and wrist extension was accompanied by finger flexion. Both the line and circle drawing were performed repetitively at four speed levels, ranging from slow to "as fast as possible" movements. The analysis of the line drawing revealed differential variability and temporal characteristics across the four movement patterns. While the equivalent pattern had characteristics of performance similar to those observed in the wrist-only and fingers-only pattern, the nonequivalent pattern was more variable and was executed slower when as fast as possible movement was required, compared to the other three patterns. The circle-drawing task also revealed intrinsic tendencies in coordination of the wrist and fingers. These tendencies were manifested by a spontaneous transition of the circular path of the pen tip to a tilted oval with increases in movement speed. The transition to the oval shape was accompanied by decreases in relative phase between the wrist and finger movements, whereas amplitudes of these movements were not affected by movement speed manipulations. The results suggest that subjects did not display a tendency to decrease the number of joints involved when executing the patterns that required simultaneous wrist and finger movements. Instead, there were preferences during these patterns to integrate wrist and finger movements with low relative phase. The findings are interpreted in terms of biomechanical constraints imposed on the wrist-finger linkage. This interpretation was further examined by testing two left-handed subjects. The data obtained showed symmetrical preferences in joint coordination. Collectively, the findings support a supposition that the shape of cursive letters may have been adjusted to the biomechanical structure of the hand to facilitate the motor act of handwriting.     

Elbow impedance during goal-directed movements

The mechanical properties and reflex actions of muscles crossing the elbow joint were examined during a 60-deg voluntary elbow extension movement. Brief unexpected torque pulses of identical magnitude and time-course (20-Nm extension switching to 20-Nm flexion within 30 ms) were introduced at various points of a movement in randomly selected trials. Single pulses were injected in different trials, some before movement onset and some either during early, mid, late or ending stages of the movement. Changes in movement trajectory induced by a torque pulse were determined over the first 50 ms by a nearest-neighbor prediction algorithm, and then a modified K-B-I (stiffness-damping-inertia) model was fit to the responses. The stiffness and damping coefficients estimated during voluntary movements were compared to values recorded during trials in which subjects were instructed to strongly co-contract while maintaining a static posture. This latter protocol was designed to help determine the maximum impedance a subject could generate. We determined that co-contraction increased joint stiffness greatly, well beyond that recorded under control conditions. In contrast, the stiffness magnitudes were quite small during routine voluntary movements, or when the subjects relaxed their limb. Furthermore, the damping coefficients were always significant and increased measurably at the end of movement. Reflex activity, as measured by EMG responses in biceps and triceps brachii, showed highly variable responses at latencies of 160 ms or greater. These reflexes tended to activate both elbow flexors and extensors simultaneously. These findings suggest that very low intrinsic muscle stiffness values recorded during point-to-point motion render an equilibrium point or impedance control approach implausible as a means to regulate movement trajectories. In particular, muscle that is shortening against inertial loads seems to exhibit much smaller stiffness than similarly active isometric muscle, although some degree of damping is always present and does not simply co-vary with stiffness. Although the limb muscles can be co-contracted statically or during movement with an observable increase in stiffness and even task performance, this control strategy is rarely utilized, presumably due to the greater energetic cost.Part of this work was presented earlier in abstract form (Popescu and Rymer 1999)