The role of intersegmental dynamics during rapid limb oscillations

The interactive dynamic effects of muscular, inertial and gravitational moments on rapid, multi-segmented limb oscillations were studied. Using three-segment, rigid-body equations of motion, hip, knee and ankle intersegmental dynamics were calculated for the steady-state cycles of the paw-shake response in adult spinal cats. Hindlimb trajectories were filmed to obtain segmental kinematics, and myopotentials of flexors and extensors at each of the three joints were recorded synchronously with the ciné film. The segmental oscillations that emerged during the paw-shake response were a consequence of an interplay between active and passive musculotendinous forces, inertial forces, and gravity. During steady-state oscillations, the amplitudes of joint excursions, peak angular velocities, and peak angular accelerations increased monotonically and significantly in magnitude from the proximal joint (hip) to the most distal joint (ankle). In contrast to these kinematic relationships, the maximal values of net moments at the hip and knee were equal in magnitude, but of significantly lower magnitude than the large net moment at the ankle joint. At both the ankle and the knee, the flexor and extensor muscle moments were equal, but at the hip the magnitude of the peak flexor muscle moment was significantly greater than the extensor muscle moment. Muscle moments at the hip not only acted to counterbalance accelerations of the more distal segments, but also acted to maintain the postural orientation of the hindlimb. Large muscle moments at the knee functioned to counterbalance the large inertial moments generated by the large angular accelerations of the paw. At the ankle, the muscle moments dominated the generation of the paw accelerations. At the ankle and the knee, muscle moments controlled limb dynamics by slowing and reversing joint motions, and the active muscle forces contributing to ankle and knee moments were derived from lengthening of active musculotendinous units. In contrast to the more distal joints, the active muscles crossing the hip predominantly shortened as a result of the interplay among inertial forces and gravitational moments. The muscle function and kinetic data explain key features of the complex interactions that occur between central control mechanisms and multi-segmented, oscillating limb segments during the paw-shake response.     

Simplified dynamics model of planar two-joint arm movements

A theoretical framework is presented that describes a way in which the inverse dynamics equations of motion of planar two-joint arm movements (EX-model) are reformulated in a simple form. A single point was assumed to define both the wrist and elbow joint centers, and thus the motion of two points in extrinsic space was represented by second-order differential equations to provide the variables in the reformulation (RE-) model. Through an analytical processes, it was shown that the RE-model for reproducing the shoulder joint torque consists of the linearly scaled moment per unit mass responsible for accelerating the wrist and elbow points about the shoulder joint, while that for reproducing the elbow joint torque consists of the linearly scaled moment per unit mass responsible for accelerating the wrist point about the elbow. The scaling factors for variables in the RE-model were based solely on the values for segment lengths, while in the EX-model the inertial parameter data for the segments are involved in its representation. The inertial parameter data of six-arm specimens from the cadaver experiment of Chandler et al. (1975, AMRL Technical Report, Wright-Patterson Air Force Base, OH) were used to develop and verify the numeric solutions of the RE-model. The adequacy of the model varied somewhat among subjects, but minor changes of the physical parameters of the arm segments enabled perfect reformulation, regardless of the specimens. The potential abilities of the RE-model to deal with the complexities in motor control with more simple control schemes are discussed.     

Kinematic invariants during cyclical arm movements

It has been observed that the motion of the arm end-point (the hand, fingertip or the tip of a pen) is characterized by a number of regularities (kinematic invariants). Trajectory is usually straight, and the velocity profile has a bell shape during point-to-point movements. During drawing movements, a two-thirds power law predicts the dependence of the end-point velocity on the trajectory curvature. Although various principles of movement organization have been discussed as possible origins of these kinematic invariants, the nature of these movement trajectory characteristics remains an open question. A kinematic model of cyclical arm movements derived in the present study analytically demonstrates that all three kinematic invariants can be predicted from a two-joint approximation of the kinematic structure of the arm and from sinusoidal joint motions. With this approach, explicit expressions for two kinematic invariants, the two-thirds power law during drawing movements and the velocity profile during point-to-point movements are obtained as functions of arm segment lengths and joint motion parameters. Additionally, less recognized kinematic invariants are also derived from the model. The obtained analytical expressions are further validated with experimental data. The high accuracy of the predictions confirms practical utility of the model, showing that the model is relevant to human performance over a wide range of movements. The results create a basis for the consolidation of various existing interpretations of kinematic invariants. In particular, optimal control is discussed as a plausible source of invariant characteristics of joint motions and movement trajectories.     

Long-latency reflexes of the human arm reflect an internal model of limb dynamics

A key feature of successful motor control is the ability to counter unexpected perturbations. This process is complicated in multijoint systems, like the human arm, by the fact that loads applied at one joint will create motion at other joints [1-3]. Here, we test whether our most rapid corrections, i.e., reflexes, address this complexity through an internal model of the limb's mechanical properties. By selectively applying torque perturbations to the subject's shoulder and/or elbow, we revealed a qualitative difference between the arm's short-latency/spinal reflexes and long-latency/cortical reflexes. Short-latency reflexes of shoulder muscles were linked exclusively to shoulder motion, whereas its long-latency reflexes were sensitive to both shoulder and elbow motion, i.e., matching the underlying shoulder torque. In fact, a long-latency reflex could be evoked without even stretching or lengthening the shoulder muscle but by displacing just the elbow joint. Further, the shoulder's long-latency reflexes were appropriately modified across the workspace to account for limb-geometry changes that affect the transformation between joint torque and joint motion. These results provide clear evidence that long-latency reflexes possess an internal model of limb dynamics, a degree of motor intelligence previously reserved for voluntary motor control [3-5]. The use of internal models for both voluntary and reflex control is consistent with substantial overlap in their neural substrates and current notions of intelligent feedback control [6-8].     

Dynamic interactions between limb segments during planar arm movement

Movement of multiple segment limbs requires generation of appropriate joint torques which include terms arising from dynamic interactions among the moving segments as well as from such external forces as gravity. The interaction torques, arising from inertial, centripetal, and Coriolis forces, are not present for single joint movements. The significance of the individual interaction forces during reaching movements in a horizontal plane involving only the shoulder and elbow joints has been assessed for different movement paths and movement speeds. Trajectory formation strategies which simplify the dynamics computation are presented.     

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     

A method for measuring endpoint stiffness during multi-joint arm movements

Current methods for measuring stiffness during human arm movements are either limited to one-joint motions, or lead to systematic errors. The technique presented here enables a simple, accurate and unbiased measurement of endpoint stiffness during multi-joint movements. Using a computer-controlled mechanical interface, the hand is displaced relative to a prediction of the undisturbed trajectory. Stiffness is then computed as the ratio of restoring force to displacement amplitude. Because of the accuracy of the prediction (< 1 cm error after 200 ms) and the quality of the implementation, the movement is not disrupted by the perturbation. This technique requires only 13 as many trials to identify stiffness as the method of Gomi and Kawato (1997, Biological Cybernetics 76, 163-171) and may, therefore, be used to investigate the evolution of stiffness during motor adaptation.     

Localized muscular fatigue duration, EMG parameters and accuracy of rapid limb movements

While much is known about the physiological basis of local muscular fatigue, little is known about the kinematic and electromyographic (EMG) consequences of brief fatiguing isometric contractions. Five male subjects performed a horizontal elbow flexion-extension reversal movement over 90° in 250 ms to reversal before and after one of five single maximal isometric elbow flexions ranging in duration from 15–120 s. Surface EMG signals were recorded from the biceps brachii, the long head of the triceps, the clavicular portion of the pectoralis major, and the posterior deltoid. Spatial and temporal errors were computed from potentiometer output. During the fatiguing bouts, maximum voluntary force dropped linearly an average of 4% in the 15 s condition and 58% in the 120 s condition relative to maximum force. The associated biceps rectified-integrated EMG signal increased from the onset of each fatigue bout for 15–30 s, then decreased over the remainder of the longer bouts. Following the fatigue bout, subjects undershot the target distance on the first movement trial in all conditions. Following short fatigue durations (i.e. 15–30 s), the peak biceps EMG amplitude was disrupted and movement velocity decreased, but both measures recovered within seconds. As fatigue duration increased, progressive decreases in peak velocity occurred with increased time to reversal, reduced EMG amplitude, and longer recovery times. However, the relative timing of the EMG pattern was maintained suggesting the temporal structure was not altered by fatigue. The findings suggest that even short single isometric contractions can disrupt certain elements of the motor control system.     

Changes in respiratory movements of the human vocal cords during hyperpnea

Five healthy young subjects were studied to assess the changes in vocal cord movements that occur between resting breathing and hyperpnea. Both hypercapnia and exercise induced decreases in the extent of narrowing of the glottic aperture occurring during expiration. In addition, four of the subjects showed a significant positive rank correlation between the extent of narrowing of the glottis and the observed length of the expiratory phase of the respiratory cycle. These results indicate that the braking of expiratory airflow by movements of the vocal cords toward the midline is reduced during hyperpnea at the same time that expiratory time is decreased.     

Changes in the pericellular matrix during differentiation of limb bud mesoderm

Mesodermal cells in the developing chick embryo limb bud appear morphologically homogeneous until stage 21. At stage 22 the prechondrogenic and premyogenic areas begin to condense, culminating in the appearance of cartilage and muscle by stage 25-26. We have examined changes in the hyaluronate-dependent pericellular matrices elaborated by mesodermal cells of the limb bud from different developmental stages and the corresponding changes in production of cell surface-associated and secreted glycosaminoglycans. When placed in culture, most early mesodermal cells (stage 17 lateral plate and stage 19 limb bud) exhibited pericellular coats as visualized by the exclusion of particles. These coats were removed by treatment of the cultures with Streptomyces hyaluronidase. Cells from stage 20-21 limb buds (precondensation) had smaller coats, whereas cells derived from stage 22, 24, and 26 limb buds (condensed chondrogenic and myogenic regions) lacked coats. However, coats were reformed during subsequent cytodifferentiation of chondrocytes; chondrocytes from stage 28 and 30 limb buds, and more mature chondrocytes from stage 38 tibiae, had pericellular coats. Thus, cytodifferentiation of cartilage is accompanied by extensive intercellular matrix accumulation in vivo and reacquisition of pericellular coats in vitro. Although their structure was still dependent on hyaluronate, chondrocyte coats were associated with increased proteoglycan content compared to the coats of early mesodermal cells. The amount of incorporation of [3H]acetate into cell surface hyaluronate remained relatively constant from stages 17 to 38, whereas in the medium compartment, incorporation into hyaluronate was more than 4-fold greater by stage 17 and 19 mesodermal cells than by cells from stages between 20 and 38. However, there was a progressive increase in incorporation into cell surface and medium chondroitin sulfate throughout these developmental stages. Thus, at the time of cellular condensation in the limb bud in vivo, we have observed a reduction in size of hyaluronate-dependent pericellular coats and a dramatic change in the relative proportion of hyaluronate and chondroitin sulfate produced by the mesodermal cells in vitro.     

Changes in Bereitschaftspotential during fatiguing and non-fatiguing hand movements

This study investigates the electroencephalographic Bereitschaftspotential (Bp) during muscular-fatiguing and non-fatiguing rhythmical hand contractions at 20%, 50%, and 80% maximum voluntary contraction (MVC). A feedback arrangement was provided so that subjects were able to adjust the force as required. The results confirm that Bp depends on force level. An increase in force results in a Bp increase. Further, they show that muscular-fatiguing contractions at 80% MVC are accompanied by an increased Bp. This could be the result of an increase in the central nervous activation required when preparing for motor activity with fatigued muscles. A decrease in Bp was observed during non-fatiguing repetitive hand contractions at 50% MVC. Possibly, the decrease reflects a decrease in subjects intentional involvement due to the monotony of the exercise. Repetitive movements at 20% MVC, which require a high degree of concentration and attention to adjust exactly to this very small force level, also result in an increased Bp, perhaps due to the higher intentional involvement. It may be concluded that the influence of muscular fatigue on Bp should be investigated with consideration of the psychological aspects of repetitive movements.     

Motor control of voluntary arm movements

The motor control of pointing and reaching-to-grasp movements was investigated using two different approaches (kinematic and modelling) in order to establish whether the type of control varies according to modifications of arm kinematics. Kinematic analysis of arm movements was performed on subjects' hand trajectories directed to large and small stimuli located at two different distances. The subjects were required either to grasp and to point to each stimulus. The kinematics of the subsequent movement, during which subject's hand came back to the starting position, were also studied. For both movements, kinematic analysis was performed on hand linear trajectories as well as on joint angular trajectories of shoulder and elbow. The second approach consisted in the parametric identification of the black box (ARMAX) model of the controller driving the arm movement. Such controller is hypothesized to work for the correct execution of the motor act. The order of the controller ARMAX model was analyzed with respect to the different experimental conditions (distal task, stimulus size and distance). Results from kinematic analysis showed that target distance and size influenced kinematic parameters both of angular and linear displacements. Nevertheless, the structure of the motor program was found to remain constant with distane and distal task, while it varied with precision requirements due to stimulus size. The estimated model order of the controller confirmed the invariance of the control law with regard to movement amplitude, whereas it was sensitive to target size.     

Impaired agonists recruitment during voluntary arm movements in patients affected by spasmodic torticollis

We have investigated the electromyographic (EMG) and kinematic characteristics of horizontal arm extension movements in patients affected by idiopathic cervical dystonia (ICD) as well as in normal subjects. In spite of the lack of an overt dystonic involvement of the muscles acting at upper arm level, all these patients were considerably bradykinetic. Although the degree of bradykinesia observed was comparable to that previously reported for the body segment directly affected by this patholgy (21,15,8), the EMG analysis of the agonist muscles indicated a specific pathophysiological mechanism. In particular, the recruitment of the posterior deltoid (pD) in ICD patients was severely impaired within the initial phase (130 ms) of the movement. On the other hand, within the same time span, the activation of the mD, a muscle that plays a more important postural role than the pD, was not significantly different between patients and normal subjects. This reduced recruitment in the initial phase of the AG1 appears responsible of the slowness of voluntary movements.     

Essential role of MARCKS in cortical actin dynamics during gastrulation movements

Myristoylated alanine-rich C kinase substrate (MARCKS) is an actin-binding, membrane-associated protein expressed during Xenopus embryogenesis. We analyzed its function in cytoskeletal regulation during gastrulation. Here, we show that blockade of its function impaired morphogenetic movements, including convergent extension. MARCKS was required for control of cell morphology, motility, adhesion, protrusive activity, and cortical actin formation in embryonic cells. We also demonstrate that the noncanonical Wnt pathway promotes the formation of lamellipodia- and filopodia-like protrusions and that MARCKS is necessary for this activity. These findings show that MARCKS regulates the cortical actin formation that is requisite for dynamic morphogenetic movements.     

Compensation of large motion sensor displacements during long recordings of limb movements

In motion capture applications using electromagnetic tracking systems the process of anatomical calibration associates the technical frames of sensors attached to the skin with the human anatomy. Joint centers and axes are determined relative to these frames. A change of orientation of the sensor relative to the skin renders this calibration faulty. This sensitivity regarding sensor displacement can turn out to be a serious problem with movement recordings of several minutes duration. We propose the “dislocation distance” as a novel method to quantify sensor displacement and to detect gradual and sudden changes of sensor orientation. Furthermore a method to define a so called fixed technical frame is proposed as a robust reference frame which can adapt to a new sensor orientation on the skin. The proposed methods are applied to quantify the effects of sensor displacement of 120 upper and lower limb movement recordings of newborns revealing the need for a method to compensate for sensor displacement. The reliability of the fixed technical frame is quantified and it is shown that trend and dispersion of the dislocation distance can be significantly reduced. A working example illustrates the consequences of sensor displacement on derived angle time series and how they are avoided using the fixed technical frame.     

Cortical regions activated after rapid eye movements during REM sleep

The present study investigated the cortical regions activated during rapid eye movement (REM) sleep by identifying the sources of electric currents of brain potentials related to rapid eye movements using low-resolution brain electromagnetic tomography (LORETA). The brain potentials measured were the lambda response (P1 and P2) during wakefulness and the lambda-like response (P1r and P2r) during REM sleep. Fifteen healthy university students participated in this study. During wakefulness, the sources of the electric current of the lambda response (P1 and P2) were estimated to be in the primary and secondary visual cortices (BA 17, 18). During REM sleep, the P1r has a source in a higher order visual area (precuneus; BA 7, 31) and P2r comes from the primary and secondary visual cortices (BA 17, 18). In addition, the density of electric current in the premotor and fronto-central regions including anterior cingulate gyrus was higher after rapid eye movements, which was a discriminative feature of REM sleep. The results of this study suggest that these activities that occur after rapid eye movements might underlie the generation of vivid visual images of dreaming.

     

Comparative analysis of the kinematics of hind limb movements in rats during different kinds of locomotion

A comparative analysis of phases of the locomotor cycle and the dynamics of changes in hind limb joint angles during swimming and stepping movements (on a treadmill), involving the fore- and hind limbs to different degrees, were undertaken in rats. Differences in the sequence and degree of changes in joint angles during locomotion of the types investigated were participation of the forelimbs in locomotion was found to be accompanied by more marked forward carrying of the hind limb. Dependence of the swing phase on duration of the cycle was observed and differences were found in the period of protraction of the limb (F period) during swimming and stepping. The role of central spinal processes and influences of peripheral afferents in the formation of different types of locomotion is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 2, pp. 189–198, March–April, 1985.