Stability and motor adaptation in human arm movements

In control, stability captures the reproducibility of motions and the robustness to environmental and internal perturbations. This paper examines how stability can be evaluated in human movements, and possible mechanisms by which humans ensure stability. First, a measure of stability is introduced, which is simple to apply to human movements and corresponds to Lyapunov exponents. Its application to real data shows that it is able to distinguish effectively between stable and unstable dynamics. A computational model is then used to investigate stability in human arm movements, which takes into account motor output variability and computes the force to perform a task according to an inverse dynamics model. Simulation results suggest that even a large time delay does not affect movement stability as long as the reflex feedback is small relative to muscle elasticity. Simulations are also used to demonstrate that existing learning schemes, using a monotonic antisymmetric update law, cannot compensate for unstable dynamics. An impedance compensation algorithm is introduced to learn unstable dynamics, which produces similar adaptation responses to those found in experiments.     

Using voluntary motor commands to inhibit involuntary arm movements

A hallmark of voluntary motor control is the ability to stop an ongoing movement. Is voluntary motor inhibition a general neural mechanism that can be focused on any movement, including involuntary movements, or is it mere termination of a positive voluntary motor command? The involuntary arm lift, or ‘floating arm trick’, is a distinctive long-lasting reflex of the deltoid muscle. We investigated how a voluntary motor network inhibits this form of involuntary motor control. Transcranial magnetic stimulation of the motor cortex during the floating arm trick produced a silent period in the reflexively contracting deltoid muscle, followed by a rebound of muscle activity. This pattern suggests a persistent generator of involuntary motor commands. Instructions to bring the arm down voluntarily reduced activity of deltoid muscle. When this voluntary effort was withdrawn, the involuntary arm lift resumed. Further, voluntary motor inhibition produced a strange illusion of physical resistance to bringing the arm down, as if ongoing involuntarily generated commands were located in a ‘sensory blind-spot’, inaccessible to conscious perception. Our results suggest that voluntary motor inhibition may be a specific neural function, distinct from absence of positive voluntary motor commands.     

Effect of muscle fatigue on internal model formation and retention during reaching with the arm.

The motor system adapts to novel dynamic environments by forming internal models that predict the muscle forces needed to move skillfully. The goal of this study was to determine how muscle fatigue affects internal model formation during arm movement and whether an internal model acquired while fatigued could be recalled accurately after rest. Twelve subjects adapted to a viscous force field applied by a lightweight robot as they reached to a target. They then reached while being resisted by elastic bands until they could no longer touch the target. This protocol reduced the strength of the muscles used to resist the force field by approximately 20%. The bands were removed, and subjects adapted again to the viscous force field. Their adaptive ability, quantified by the amount and time constant of adaptation, was not significantly impaired following fatigue. The subjects then rested, recovering approximately 70% of their lost force-generation ability. When they reached in the force field again, their prediction of the force field strength was different than in a nonfatigued state. This alteration was consistent with the use of a higher level of effort than normally used to counteract the force field. These results suggest that recovery from fatigue can affect recall of an internal model, even when the fatigue did not substantially affect the motor system's ability to form the model. Recovery from fatigue apparently affects recall because the motor system represents internal models as a mapping between effort and movement and relies on practice to recalibrate this mapping.     

A fuzzy-genetic model for estimating forces from electromyographical activity of antagonistic muscles due to planar lower arm movements: the effect of nonlinear muscle properties

The aim of this paper is to create a model for mapping the surface electromyogram (EMG) signals to the force that generated by human arm muscles. Because the parameters of each person's muscle are individual, the model of the muscle must have two characteristics: (1) The model must be adjustable for each subject. (2) The relationship between the input and output of model must be affected by the force-length and the force-velocity behaviors are proven through Hill's experiments. Hill's model is a kinematic mechanistic model with three elements, i.e. one contractile component and two nonlinear spring elements.In this research, fuzzy systems are applied to improve the muscle model. The advantages of using fuzzy system are as follows: they are robust to noise, they prove an adjustable nonlinear mapping, and are able to model the uncertainties of the muscle.Three fuzzy coefficients have been added to the relationships of force-length (active and passive) and force-velocity existing in Hill's model. Then, a genetic algorithm (GA) has been used as a biological search method that can adjust the parameters of the model in order to achieve the optimal possible fit.Finally, the accuracy of the fuzzy genetic implementation Hill-based muscle model (FGIHM) is invested as following: the FGIHM results have 12.4% RMS error (in worse case) in comparison to the experimental data recorded from three healthy male subjects. Moreover, the FGIHM active force-length relationship which is the key characteristics of muscles has been compared to virtual muscle (VM) and Zajac muscle model. The sensitivity of the FGIHM has been evaluated by adding a white noise with zero mean to the input and FGIHM has proved to have lower sensitivity to input noise than the traditional Hill's muscle model.     

A model of force and impedance in human arm movements

This paper describes a simple computational model of joint torque and impedance in human arm movements that can be used to simulate three-dimensional movements of the (redundant) arm or leg and to design the control of robots and human-machine interfaces. This model, based on recent physiological findings, assumes that (1) the central nervous system learns the force and impedance to perform a task successfully in a given stable or unstable dynamic environment and (2) stiffness is linearly related to the magnitude of the joint torque and increased to compensate for environment instability. Comparison with existing data shows that this simple model is able to predict impedance geometry well.     

Comparison of an EMG-based and a stress-based method to predict shoulder muscle forces

The estimation of muscle forces in musculoskeletal shoulder models is still controversial. Two different methods are widely used to solve the indeterminacy of the system: electromyography (EMG)-based methods and stress-based methods. The goal of this work was to evaluate the influence of these two methods on the prediction of muscle forces, glenohumeral load and joint stability after total shoulder arthroplasty. An EMG-based and a stress-based method were implemented into the same musculoskeletal shoulder model. The model replicated the glenohumeral joint after total shoulder arthroplasty. It contained the scapula, the humerus, the joint prosthesis, the rotator cuff muscles supraspinatus, subscapularis and infraspinatus and the middle, anterior and posterior deltoid muscles. A movement of abduction was simulated in the plane of the scapula. The EMG-based method replicated muscular activity of experimentally measured EMG. The stress-based method minimised a cost function based on muscle stresses. We compared muscle forces, joint reaction force, articular contact pressure and translation of the humeral head. The stress-based method predicted a lower force of the rotator cuff muscles. This was partly counter-balanced by a higher force of the middle part of the deltoid muscle. As a consequence, the stress-based method predicted a lower joint load (16% reduced) and a higher superior–inferior translation of the humeral head (increased by 1.2 mm). The EMG-based method has the advantage of replicating the observed cocontraction of stabilising muscles of the rotator cuff. This method is, however, limited to available EMG measurements. The stress-based method has thus an advantage of flexibility, but may overestimate glenohumeral subluxation.     

Learning the dynamics of reaching movements results in the modification of arm impedance and long-latency perturbation responses

 Some characteristics of arm movements that humans exhibit during learning the dynamics of reaching are consistent with a theoretical framework where training results in motor commands that are gradually modified to predict and compensate for novel forces that may act on the hand. As a first approximation, the motor control system behaves as an adapting controller that learns an internal model of the dynamics of the task. It approximates inverse dynamics and predicts motor commands that are appropriate for a desired limb trajectory. However, we had previously noted that subtle motion characteristics observed during changes in task dynamics challenged this simple model and raised the possibility that adaptation also involved sensory–motor feedback pathways. These pathways reacted to sensory feedback during the course of the movement. Here we hypothesize that adaptation to dynamics might also involve a modification of how the CNS responds to sensory feedback. We tested this through experiments that quantified how the motor system's response to errors during voluntary movements changed as it adapted to dynamics of a force field. We describe a nonlinear approach that approximates the impedance of the arm, i.e., force response as a function of arm displacement trajectory. We observe that after adaptation, the impedance function changes in a way that closely matches and counters the effect of the force field. This is particularly prominent in the long-latency (>100 ms) component of response to perturbations. Therefore, it appears that practice not only modifies the internal model with which the brain generates motor commands that initiate a movement, but also the internal model with which sensory feedback is integrated with the ongoing descending commands in order to respond to error during the movement. Received: 10 January 2001 / Accepted in revised form: 30 May 2001     

Motor unit recruitment in the trapezius muscle with special reference to coarse arm movements.

Chronic shoulder pain is common in a variety of occupations. The "Cinderella hypothesis" suggests that the pain originates from damaged type I muscle fibres driven into degenerative processes as a result of too long activation and too short recovery time. The main purpose of this study was to investigate if the same motor units are active during all phases of coarse arm movements. Eight healthy volunteers participated in the study. Intramuscular electromyographic signals were picked up with a four-lead fine wire electrode, during a unilateral straight arm movement. The movement started with either (part 1) an abduction or a flexion, then (2) a movement in the horizontal plane from the sagittal to the frontal plane or vice versa, and finally (3) an adduction or an extension to the start position. The movement cycle was performed in three different speeds, slow, medium, and high, with one, two or five cycles per 20 s, respectively. On an average, the motor unit action potentials (MUAPs) of 6 motor units (range, 1-15) were identified per trail. In total 94% of the MUAP trains that were identified showed firings in all 3 parts of the movements. The findings support the Cinderella hypothesis, although there is a need to further investigate the temporal pattern of long-term motor unit activity.     

Endogenous patterns of photomechanical movements in teleosts and their relation to activity rhythms

Summary The retinal rods, cones and epithelial pigment of most lower vertebrates display rhythmic photomechanical (retinomotor) migrations in response to changes in ambient lighting conditions. This study examines the extent of these migrations in the absence of the daily changes in illumination (constant darkness and constant light) in three species of teleosts. Salmo trutta, a crepuscularly active fish, showed two peaks of light adaptation occurring around dawn and dusk when kept in constant darkness. Tinca tinca, a nocturnal species, also showed an endogenous rhythm during extended periods of darkness, but, unlike Salmo trutta, it was light-adapted throughout what would normally have been day. At the maximal extent of migration under conditions of continual darkness, the pigment migrated 59% as much as it did during a normal light/dark cycle. Nannacara anomala, a tropical diurnally active species, showed a similar but more pronounced rhythm than Tinea tinea for all 3 days of experimental darkness, behaving essentially identically to fish exposed to a light/dark cycle. Nannacara anomala also displayed a weak rhythm when kept in constant light.It is concluded from these and previous results that the pattern of endogenous photomechanical movement depends both on the activity pattern of a species and on the constancy of the lighting conditions to which it has been exposed during its lifetime.     

Summation of motor unit forces in rat medial gastrocnemius muscle

The summation of contractile forces of motor units (MUs) was analyzed by comparing the recorded force during parallel stimulation of two and four individual MUs or four groups of MUs to the algebraic sum of their individual forces. Contractions of functionally-isolated single MUs of the medial gastrocnemius muscle were evoked by electrical stimulation of thin filaments of the split L5 or L4 ventral roots of spinal nerves. Additionally, contractions of large groups of MUs were evoked by stimuli delivered to four parts of the divided L5 ventral root. Single twitches, 40 Hz unfused tetani, and 150 Hz fused maximum tetani were recorded. In these experimental situations the summation was more effective for unfused tetani than for twitches or maximum tetani. The results obtained for pairs of MUs were highly variable (more- or less-than-linear summation), but coactivation of more units led to progressively weaker effects of summation, which were usually less-than-linear in comparison to the algebraic sums of the individual forces. The variability of the results highlights the importance of the structure of the muscle and the architecture of its MUs. Moreover, the simultaneous activity of fast and slow MUs was considerably more effective than that of two fast units.     

An EMG-level muscle model for a fast arm movement to target

A model of human muscle action is presented for a maximally fast, large-amplitude forearm movement to target. the inputs to the model are approximately the biceps and triceps EMG envelopes over a single movement. The model's output gives the corresponding displacement angle of the forearm about a fixed elbow position as a function of time. The idea of the model is to conceive of both EMG input drives as successions of millisecond input pulses, with each pulse resulting in a muscle tension twitch. Every twitch is amplitude-scaled, parametrically-shaped, and duration-limited as a function of the muscle's contractile history thus far in the movement. The muscle tension at any time t is the sum of the residual tension levels of all twitches begun before t. The model was developed and tested with special reference to two subjects: one, according to the model dynamics, was a comparatively slow-twitch type, and the other modelled as a fast-twitch type. Good agreement was found between model output and subject response data whenever the subject's EMG's were synchronous. The model can be used to characterize each subject's responses by a suite of twitch characteristics. This will enable us to check the accepted but now suspect correlation between muscle biopsy-and performance-determined muscle twitch type.This study was supported by contract DAMD 17-80-C-0101 from the U.S. Army Medical Research and Development Command. The views, opinions and/or findings contained in this report are those of the authors and should not be construed as an official Department of the Army position, policy, or decision, unless so designated by other documentation     

Na activation delays and their relation to inactivation in frog skeletal muscle

Summary Delays in the development of activation of Na currents were studied using voltage-clamped frog skeletal muscle fibers. Na currents elicited by a depolarizing voltage step from a hyperpolarized membrane potential were delayed in their activation when compared to Na currents elicited from the resting potential. The magnitude of the delay increased with larger hyperpolarizing potentials and decreased with larger depolarizing test potentials. Delays in activation observed following chloramine-T treatment that partially removes inactivation did not differ from delays observed before treatment. Longer exposures of the muscle fiber to chloramine-T led to a complete loss of inactivation, coincident with an elimination of the hyperpolarization-induced delays in activation. Steady-state slow inactivation was virtually unaffected by prolonged exposures of the fibers to chloramine-T that eliminated fast inactivation. The results show that chloramine-T acts at a number of sites to alter both activation and inactivation. Markov model simulations of the results show that chloramine-T alters fundamental time constants of the system by altering both activation and inactivation rate constants.     

Long-lasting inhibition of the soleus muscle H reflex related to realization of voluntary arm movements in humans

In healthy humans, we recorded the H reflex induced by transcutaneous stimulation of the tibial nerve (recording from the soleus muscle). In subjects in the lying position, we studied changes in the H reflex values after preceding voluntary arm movements realized with a maximum velocity after presentation of an acoustic signal. On the 200th to 300th msec after forearm flexion, long-lasting inhibition of the H reflex developed following a period of initial facilitation and reached the maximum, on average, 700 msec from the moment of the movement. Flexion of the contralateral upper limb in the elbow joint induced deeper inhibition than analogous movement of the ipsilateral arm. Long-lasting clear inhibition of the H reflex developed after arm flexion in the elbow joint but was slightly expressed after finger clenching. After inhibition reached the maximum, its time course was satisfactorily approximated by a logarithmic function of the time interval between the beginning of the conditioning voluntary movement and presentation of the test stimulus. Durations of inhibition calculated using a regression equation were equal to 6.6 sec and 8.5 sec after ipsilateral and contralateral elbow-joint flexions, respectively. Inhibition was not eliminated under conditions of tonic excitation of motoneurons of the tested muscle upon voluntary foot flexion. Long-lasting inhibition of the H reflex was also observed after electrical stimulation-induced flexions of the upper limb. The obtained data indicate that movements of the upper limb cause reflex long-lasting presynaptic inhibition of the soleus-muscle H reflex that can play a noticeable role in redistribution of the muscle tone during motor activity. Neirofiziologiya/Neurophysiology, Vol. 40, No. 3, pp. 221–227, May–June, 2008.     

A proprioception based regulation model to estimate the trunk muscle forces

Evaluation of loads acting on the spine requires the knowledge of the muscular forces acting on it, but muscles redundancy necessitates developing a muscle forces attribution strategy. Optimisation, EMG, or hybrid models allow evaluating muscle force patterns, yielding a unique muscular arrangement or/and requiring EMG data collection. This paper presents a regulation model of the trunk muscles based on a proprioception hypothesis, which searches to avoid the spinal joint overloading. The model is also compared to other existing models for evaluation. Compared to an optimisation model, the proposed alternative muscle pattern yielded a significant spine postero-anterior shear decrease. Compared to a model based on combination of optimisation criteria, present model better fits muscle activation observed using EMG (38% improvement). Such results suggest that the proposed model, based on regulation of all spinal components, may be more relevant from a physiologic point of view.     

A validated model of passive skeletal muscle to predict force and intramuscular pressure

The passive properties of skeletal muscle are often overlooked in muscle studies, yet they play a key role in tissue function in vivo. Studies analyzing and modeling muscle passive properties, while not uncommon, have never investigated the role of fluid content within the tissue. Additionally, intramuscular pressure (IMP) has been shown to correlate with muscle force in vivo and could be used to predict muscle force in the clinic. In this study, a novel model of skeletal muscle was developed and validated to predict both muscle stress and IMP under passive conditions for the New Zealand White Rabbit tibialis anterior. This model is the first to include fluid content within the tissue and uses whole muscle geometry. A nonlinear optimization scheme was highly effective at fitting model stress output to experimental stress data (normalized mean square error or NMSE fit value of 0.993) and validation showed very good agreement to experimental data (NMSE fit values of 0.955 and 0.860 for IMP and stress, respectively). While future work to include muscle activation would broaden the physiological application of this model, the passive implementation could be used to guide surgeries where passive muscle is stretched.     

Reproduction of reaching movements to memorized targets in the lack of visual control

The objective of this study was to examine the precision of reaching movements to remembered target distances in the lack of visual information. Subjects were professional basketball players and nonathlete university students. The basketball players, having well-trained manual skills, performed better than the non-athlete students. Increase in the overestimation of the remembered target distances was found under the effect of fatigue. A weight load on the sliding handle caused some decrease in the errors of reaching the remembered targets. No significant gender differences were found.     

Modulation of the soleus muscle H reflex caused by ballistic voluntary arm movements in humans

In healthy humans, we studied the influence of conditioning voluntary arm movements on the H reflex induced by transcutaneous stimulation of the tibial nerve and recorded from the soleus muscle. We examined the effects of flexion and extension of the forearm, as well as of finger clenching performed with the maximum rate. Conditioning arm movements were self-induced or realized upon presentation of a visual signal (light flash). We found that the pattern of changes in the H reflex is determined by the position of the subject’s body in the course of tests. The ipsilateral arm flexion in the elbow joint in the standing position resulted in depression of the H reflex lasting about 100 msec from the beginning of the movement, while the effect observed in the lying position (on the couch with the feet hanging free in the air) looked like a facilitation of the reflex lasting about 100 to 200 msec. The direction and dynamics of modifications of the H reflex under conditions of the use of different conditioning movements (forearm flexions/extensions and finger clenching of the ipsilateral arm, as well as contralateral forearm flexions in the elbow joint) were rather similar. We also showed that the observed facilitation of the H reflex began earlier than the voluntary arm movement (40 to 50 msec prior to the beginning). We hypothesize that these conditioning influences result from the action of central motor commands and represent the factor related to anticipatory postural rearrangements. Such rearrangements are directed toward the maintenance of equilibrium of the body in the course of a future movement. These commands depend significantly on the spatial position of the subject’s body. Neirofiziologiya/Neurophysiology, Vol. 40, No. 2, pp. 147–154, March–April, 2008.