Analysis of kinematic invariances of multijoint reaching movement

 There is a no unique relationship between the trajectory of the hand, represented in cartesian or extrinsic space, and its trajectory in joint angle or intrinsic space in the general condition of joint redundancy. The goal of this work is to analyze the relation between planning the trajectory of a multijoint movement in these two coordinate systems. We show that the cartesian trajectory can be planned based on the task parameters (target coordinates, etc.) prior to and independently of angular trajectories. Angular time profiles are calculated from the cartesian trajectory to serve as a basis for muscle control commands. A unified differential equation that allows planning trajectories in cartesian and angular spaces simultaneously is proposed. Due to joint redundancy, each cartesian trajectory corresponds to a family of angular trajectories which can account for the substantial variability of the latter. A set of strategies for multijoint motor control following from this model is considered; one of them coincides with the frog wiping reflex model and resolves the kinematic inverse problem without inversion. The model trajectories exhibit certain properties observed in human multijoint reaching movements such as movement equifinality, straight end-point paths, bell-shaped tangential velocity profiles, speed-sensitive and speed-insensitive movement strategies, peculiarities of the response to double-step targets, and variations of angular trajectory without variations of the limb end-point trajectory in cartesian space. In humans, those properties are almost independent of limb configuration, target location, movement duration, and load. In the model, these properties are invariant to an affine transform of cartesian space. This implies that these properties are not a special goal of the motor control system but emerge from movement kinematics that reflect limb geometry, dynamics, and elementary principles of motor control used in planning. All the results are given analytically and, in order to compare the model with experimental results, by computer simulations. Received: 6 April 1994/Accepted in revised form: 25 April 1995     

Analysis of an optimal control model of multi-joint arm movements

 In this paper, we propose a model of biological motor control for generation of goal-directed multi-joint arm movements, and study the formation of muscle control inputs and invariant kinematic features of movements. The model has a hierarchical structure that can determine the control inputs for a set of redundant muscles without any inverse computation. Calculation of motor commands is divided into two stages, each of which performs a transformation of motor commands from one coordinate system to another. At the first level, a central controller in the brain accepts instructions from higher centers, which represent the motor goal in the Cartesian space. The controller computes joint equilibrium trajectories and excitation signals according to a minimum effort criterion. At the second level, a neural network in the spinal cord translates the excitation signals and equilibrium trajectories into control commands to three pairs of antagonist muscles which are redundant for a two-joint arm. No inverse computation is required in the determination of individual muscle commands. The minimum effort controller can produce arm movements whose dynamic and kinematic features are similar to those of voluntary arm movements. For fast movements, the hand approaches a target position along a near-straight path with a smooth bell-shaped velocity. The equilibrium trajectories in X and Y show an ‘N’ shape, but the end-point equilibrium path zigzags around the hand path. Joint movements are not always smooth. Joint reversal is found in movements in some directions. The excitation signals have a triphasic (or biphasic) pulse pattern, which leads to stereotyped triphasic (or biphasic) bursts in muscle control inputs, and a dynamically modulated joint stiffness. There is a fixed sequence of muscle activation from proximal muscles to distal muscles. The order is preserved in all movements. For slow movements, it is shown that a constant joint stiffness is necessary to produce a smooth movement with a bell-shaped velocity. Scaled movements can be reproduced by varying the constraints on the maximal level of excitation signals according to the speed of movement. When the inertial parameters of the arm are altered, movement trajectories can be kept invariant by adjusting the pulse height values, showing the ability to adapt to load changes. These results agree with a wide range of experimental observations on human voluntary movements. Received: 4 December 1995 / Accepted in revised form: 17 September 1996     

Assessing Movement Factors in Upper Limb Kinematics Decoding from EEG Signals

The past decades have seen the rapid development of upper limb kinematics decoding techniques by performing intracortical recordings of brain signals. However, the use of non-invasive approaches to perform similar decoding procedures is still in its early stages. Recent studies show that there is a correlation between electroencephalographic (EEG) signals and hand-reaching kinematic parameters. From these studies, it could be concluded that the accuracy of upper limb kinematics decoding depends, at least partially, on the characteristics of the performed movement. In this paper, we have studied upper limb movements with different speeds and trajectories in a controlled environment to analyze the influence of movement variability in the decoding performance. To that end, low frequency components of the EEG signals have been decoded with linear models to obtain the position of the volunteer’s hand during performed trajectories grasping the end effector of a planar manipulandum. The results confirm that it is possible to obtain kinematic information from low frequency EEG signals and show that decoding performance is significantly influenced by movement variability and tracking accuracy as continuous and slower movements improve the accuracy of the decoder. This is a key factor that should be taken into account in future experimental designs.     

The discontinuous nature of motor execution II. Merging discrete and rhythmic movements in a single-joint system -- the phase entrainment effect

 Initiation of rapid discrete flexion movements is significantly altered when a secondary rhythmic movement is performed simultaneously with the same limb; the onset of a stimulus-evoked discrete movement tends to occur time-locked to the oscillation: i.e., the rhythmic movement entrains the discrete response. This nonlinear interaction may reflect a specific principle of coordination of motor tasks which are simultaneously executed with the same effector. This part II of a tripartite research report on such single-muscle multiple-task coordination investigates the contribution of the dynamic properties of the muscle and its reflex circuitry to phase entrainment. Assuming a simple threshold-linear relationship between the control signals generated by the central nervous system and the observable kinematic and electromyographic signals, a secondary rhythmic movement will cause an additional phase-dependent delay between the central “go” command and the first observable change in actual kinematics of the compound movement. Several indicators for such threshold-linear interaction are derived and tested on real data obtained in psychophysical experiments. Four healthy subjects performed rapid lateral abductions of the index finger in response to a visual “go” signal. During a portion of the experiments, subjects produced additional low-amplitude oscillatory movements before stimulus presentation with either the same finger (one-handed task), or with the index finger of the other hand (two-handed task). Results showed phase entrainment and modulation of reaction times when the cyclic and the discrete movements were simultaneously executed by the same finger. But there was no entrainment in the bimanual execution of the tasks. The model was capable of reproducing the observed effects. It is concluded that coordination of voluntary movements which are concurrently performed by the same effector involves specific discontinuous operations, which represents an essential part of the mechanism of motor coordination. Phase entrainment reflects this characteristic discontinuous behavior of the lower stages of motor execution and does not necessarily require nonlinear interaction of motor commands at higher levels of motor processing. Received: 5 September 2001 / Accepted in revised form: 19 December 2001     

Optimal trajectory formation of constrained human arm reaching movements

Opening a door, turning a steering wheel, and rotating a coffee mill are typical examples of human movements that are constrained by the physical environment. The constraints decrease the mobility of the human arm and lead to redundancy in the distribution of actuator forces (either joint torques or muscle forces). Due to this actuator redundancy, there is an infinite number of ways to form a specific arm trajectory. However, humans form trajectories in a unique way. How do humans resolve the redundancy of the constrained motions and specify the hand trajectory? To investigate this problem, we examine human arm movements in a crank-rotation task. To explain the trajectory formation in constrained point-to-point motions, we propose a combined criterion minimizing the hand contact force change and the actuating force change over the course of movement. Our experiments show a close matching between predicted and experimental data.     

Different fusimotor reflexes from the ipsi- and contralateral hind limbs of the cat assessed in the same primary muscle spindle afferents

Experiments were performed in forty-five cats anaesthetized with alpha-chloralose. The aim of the study was to investigate a sample of primary muscle spindle afferents from triceps muscle with respect to their fusimotor reflex control from ipsi- as well as contralateral hind limb. Primary muscle spindle afferents of the triceps surae muscle were recorded from the mean rate of firing and the modulation of the afferent response to sinusoidal stretching of the triceps surae muscle was determined. Test measurements were made during tonic stretch of the ipsilateral PBSt, contralateral PBSt, contralateral triceps muscle or during extension of the intact contralateral hind limb. Control measurements were made with ipsi- and contralateral PBSt as well as contralateral triceps muscles relaxed and with contralateral hind limb in resting position. The occurrence and types of fusimotor effects were assessed by comparing test to control responses. The main finding of the present investigation was the great variability in type and size of the fusimotor effects evoked by different ipsi- and contralateral reflex stimuli. Both ipsi- and contralateral stimulations gave rise to predominantly dynamic, predominantly static or mixed static and dynamic fusimotor reflexes. In the same preparation, a given reflex stimulus often caused different reflex responses in different triceps surae primary spindle afferents. In the same afferent unit, different reflex stimuli usually produced fusimotor effects which differed from each other in type and/or size. In general, contralateral whole limb extension and stretch of contralateral PBSt muscles were more potent as reflex stimuli than stretch of the ipsilateral PBSt muscle. Stretch of the contralateral triceps surae muscle was, but for a few afferent units, ineffective as reflexogenic stimulus. It is concluded that the individualized receptive profiles of the primary muscle spindle afferents, which have been postulated in earlier investigations where the effects of different stimuli have been investigated on different cell populations, still seems to hold good when the stimuli are tested on the same units. The individuality of the receptive profiles of gamma-motoneurones is discussed in relation to different motor control hypotheses.     

Fusimotor reflexes influencing secondary muscle spindle afferents from flexor and extensor muscles in the hind limb of the cat.

The purpose of this study was to investigate secondary muscle spindle afferents from the triceps-plantaris (GS) and posterior biceps and semitendinosus (PBSt) muscles with respect to their fusimotor reflex control from different types of peripheral nerves and receptors. The activity of single secondary muscle spindle afferents was recorded from dissected and cut dorsal root filaments in alpha-chloralose anaesthetized cats. Both single spindle afferents and sets of simultaneously recorded units (2-3) were investigated. The modulation and mean rate of firing of the afferent response to sinusoidal stretching of the GS and PBSts muscle were determined. Control measurements were performed in the absence of any reflex stimulation, while test measurements were made during reflex stimulation. The reflex stimuli consisted of manually performed movements of the contralateral hind limb, muscle stretches, ligament tractions and electrical stimulations of cutaneous afferents. Altogether 21 secondary spindle afferents were investigated and 20 different reflex stimuli were employed. The general responsiveness (i.e. number of significant reflex effects/number of control-test series) was 52.4%, but a considerable variation between different stimuli was found, with the highest (89.9%) for contralateral whole limb extension and the lowest (25.0%) for stretch of the contralateral GS muscle. The size of the response to a given stimulus varied considerably between different afferents, and, in the same afferent, different reflex stimuli produced effects of varying size. Most responses were characterized by an increase in mean rate of discharge combined with a decrease in modulation, indicative of static fusimotor drive (Cussons et al., 1977). Since the secondary muscle spindle afferents are part of a positive feedback loop, projecting back to both static and dynamic fusimotor neurones (Appelberg Et al., 1892 a, 1983 b; Appelberg et al., 1986), it is suggested that the activity in the loop may work like an amplified which, during some circumstances, enhance the effect of other reflex inputs to the system (Johansson et al., 1991 b).     

Modulatory multiplicity in the functional repertoire of the feeding mechanism in cichlid fishes. I. Piscivores

Among piscivorous cichlids consistent differences have been recorded between ambush and pursuit hunters with respect to electromyographic, kinematic, pressure and behavioral profiles during prey capture by high speed inertial suction. Piscivorous cichlids possess a repertoire of at least two patterns of prey capture, each of which is characterized by an extreme regularity of the kinematic, pressure, electromyographic and behavioral profiles. The nature and locomotory behavior of the prey, visually analyzed by the predator during the prestrike stalk, determine which of the two preprogrammed patterns is recruited. Agile and elusive prey invariably will elicit a preprogrammed motor output (stereotyped motor pattern) that produces the greatest suction velocities in both ambush and pursuit hunters. The greater the kinematic and suction velocities, the greater the overlap of the firing sequences of antagonistic muscle complexes. The opercular and branchiostegal apparati function as an exceedingly effective anti-backwash device, damping potential fluid oscillations within the oropharynx. Mastication occurs by triphasic movements and actions of muscles of the upper and lower pharyngeal jaws in both ambush and pursuit hunters. The lower pharyngeal jaw is acted upon by a force couple of which the fourth levator externus on one hand and the pharyngocleithralis externus and pharyngohyoideus on the other hand are the antagonistic components. Furthermore, the lower pharyngeal jaw is suspended by a muscular sling, the tension of which can be modified continuously. It is postulated that the switch from insectivorous to piscivorous feeding regimes (and perhaps vice versa) is accomplished by very minor structural and functional modifications, because the modulatory multiplicity and total range of repertories of the feeding machinery of the two trophic groups overlap significantly. Piscivorous cichlids may not have arisen by orthoselection in gradually-changing lineages, but represent the differential success of subsets from a random pool of speciation events. Adaptive features identified as characteristic for piscivory could have evolved in multiple and independent lineages at a punctuational mode and tempo.     

Modelling the maximum voluntary joint torque/angular velocity relationship in human movement

The force exerted by a muscle is a function of the activation level and the maximum (tetanic) muscle force. In "maximum" voluntary knee extensions muscle activation is lower for eccentric muscle velocities than for concentric velocities. The aim of this study was to model this "differential activation" in order to calculate the maximum voluntary knee extensor torque as a function of knee angular velocity. Torque data were collected on two subjects during maximal eccentric-concentric knee extensions using an isovelocity dynamometer with crank angular velocities ranging from 50 to 450 degrees s(-1). The theoretical tetanic torque/angular velocity relationship was modelled using a four parameter function comprising two rectangular hyperbolas while the activation/angular velocity relationship was modelled using a three parameter function that rose from submaximal activation for eccentric velocities to full activation for high concentric velocities. The product of these two functions gave a seven parameter function which was fitted to the joint torque/angular velocity data, giving unbiased root mean square differences of 1.9% and 3.3% of the maximum torques achieved. Differential activation accounts for the non-hyperbolic behaviour of the torque/angular velocity data for low concentric velocities. The maximum voluntary knee extensor torque that can be exerted may be modelled accurately as the product of functions defining the maximum torque and the maximum voluntary activation level. Failure to include differential activation considerations when modelling maximal movements will lead to errors in the estimation of joint torque in the eccentric phase and low velocity concentric phase.     

Three-dimensional kinematic analysis of influence of hand orientation and joint limits on the control of arm postures and movements

The problems related to kinematic redundancy in both task and joint space were investigated for arm prehension movements in this paper. After a detailed analysis of kinematic redundancy of the arm, it is shown that the redundancy problem is ill posed only for the control of hand orientation. An experiment was then designed to investigate the influence of hand orientation on the control of arm movements. Since movements must be made within the limits of the joints, the influence of these limits was also analyzed quantitatively. The results of the experiment confirm that the increase of movement time because of the change of object orientation is due to the lengthening of the deceleration phase disproportionately to the rest of the movement. The variation of hand path due to the change of object orientation was observed as being surprisingly small for some subjects as opposed to the large range of object orientation, implying that hand path and hand orientation could be controlled separately, thus simplifying the computational problem of inverse kinematics. Moreover, the observations from the present experiment strongly suggest that a functional segmentation of the proximal and distal joints exists and that the control of wrist motion is dissociated from the rest of joint motions. The contribution of each joint in the control of arm movements could be determined through the principle of minimum energy and minimum discomfort under the constraints of the joint limits. A simplified inverse kinematics model was tested. It shows that these hypotheses can be easily implemented in a geometric algorithm and be used to predict arm prehension postures reasonably well under the constraints of joint limits. Received: 6 August 1998 / Accepted in revised form: 16 December 1998     

Movement mechanics as a determinate of muscle structure, recruitment and coordination

During muscle contractions, the muscle fascicles may shorten at a rate different from the muscle-tendon unit, and the ratio of these velocities is its gearing. Appropriate gearing allows fascicles to reduce their shortening velocities and allows them to operate at effective shortening velocities across a range of movements. Gearing of the muscle fascicles within the muscle belly is the result of rotations of the fascicles and bulging of the belly. Variable gearing can also occur as a result of tendon length changes that can be caused by changes in the relative timing of muscle activity for different mechanical tasks. Recruitment patterns of slow and fast fibres are crucial for achieving optimal muscle performance, and coordination between muscles is related to whole limb performance. Poor coordination leads to inefficiencies and loss of power, and optimal coordination is required for high power outputs and high mechanical efficiencies from the limb. This paper summarizes key studies in these areas of neuromuscular mechanics and results from studies where we have tested these phenomena on a cycle ergometer are presented to highlight novel insights. The studies show how muscle structure and neural activation interact to generate smooth and effective motion of the body.     

A numerical simulation of muscle spindle ensemble encoding during planar movement of the human arm

We address the issue of what proprioceptive information, regarding movement of the human arm, may be provided to the central nervous system by proprioceptors located within muscles of this limb. To accomplish this we developed a numerical simulation which could provide estimates of the length regimes experienced by a set of model receptors located within some of the principal muscles of the human arm during planar movement of this limb. These receptors were assumed to have characteristics analogous to those associated with a simple model of muscle spindle signalling of movement. To this end each spindle had proprioceptive ‘channels’ associated with it. These corresponded to primary and secondary spindle afferent fibers which could provide independent afferent output regarding the parent muscle the spindle monitored. The angles of the shoulder and elbow joints attained by subjects performing a task requiring movement of the right arm in a horizontal plane to a static visual target were recorded. For this angular data the lengths and rates of change of lengths experienced by muscle fascicles, and hence the model spindles, during movement were calculated by means of the numerical simulation. The discharge rates of the simulated spindles during the movement were calculated to derive a measure of the depth of modulation, induced by the movement, for each spindle. These values were then summed for all spindles to provide a first-order approximation of spindle ensemble coding of the movement. Significant correlations (0.0001, Spearman's rank order) were found between the resulting ensemble encodings and, in order of significance, the angular velocity of the shoulder joint (), the tangential velocity of the hand (), and the angular velocity of the elbow joint (). Correlations between the angular positions of the shoulder () and elbow () were lower. These findings indicate that the ensemble profiles of the simulated muscle spindles, encode information regarding kinematic parameters of movements related to both intrinsic and extrinsic coordinate systems. This suggests that motor structures capable of deriving such an ensemble encoding would be in a position to perform the sensory-motor transformations between intrinsic and extrinsic frames of reference necessary for controlling movements planned in extrinsic coordinates. Received: 12 August 1994 / Accepted in revised form: 17 June 1996     

The role of the plant properties in point-to-point arm movements: a robustness study

In recent papers we demonstrated by means of a modeling study that the smoothness of hand paths and the bell-shaped character of hand velocity profiles which have been experimentally observed in point-to-point arm movements can be largely attributed to the biomechanical properties of the arm rather than to specific planning by the central nervous system. In this paper we present a study of the robustness of our earlier results comprising two goals: (i) the determination of the range of model parameters for which such observations remain valid, (ii) the identification of possible relationships between model parameters and kinematic variables. The results of this study imply three conclusions: (i) the valid range of the tested model parameters (namely the main muscle parameters) is large, (ii) the modeled phenomena are well behaved in that parametric changes do not give rise to bifurcations or other behavioral discontinuities in the analyzed ranges, (iii) there exist precise relationships between certain muscle parameters and the time course of the hand velocity. These results point out that the phenomena observed in our previous work are indeed robust and can lead to useful insights into the mechanisms comprising the regulatory action of the central nervous system as well as into the design principles for biologically inspired artificial arms. Received: 4 December 1995 / Accepted in revised form: 6 November 1996     

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.     

Changes in the H Reflex Preceding Voluntary Movements of the Contralateral Lower Limb in Humans

In electromyographic studies on healthy subjects, we recorded the H reflex from the right m. soleus and measured changes in the magnitude of this reflex response related to voluntary movements of the contralateral lower limb performed according to a visual signal. The effects of back and plantar flexions of the contralateral foot of the tested subject in the lying and standing positions were examined. Changes in the H reflex magnitude began to be recorded 60 to 90 msec prior to voluntary movements of the contralateral limb. When the subject was in the lying position, these changes looked like facilitation of the H reflex at both types of movement of the contralateral foot. When the subject stood, facilitation preceded back flexion of the foot of this extremity, while plantar flexion was preceded by inhibition of the tested H reflex. Our results show that the pattern of preliminary changes in the muscle tone of one of the lower limbs is determined by the type of future movement of another limb and peculiarities of the support function realized by this limb.     

A neural network model for limb trajectory formation

This paper deals with the problem of representing and generating unconstrained aiming movements of a limb by means of a neural network architecture. The network produced time trajectories of a limb from a starting posture toward targets specified by sensory stimuli. Thus the network performed a sensory-motor transformation. The experimenters trained the network using a bell-shaped velocity profile on the trajectories. This type of profile is characteristic of most movements performed by biological systems. We investigated the generalization capabilities of the network as well as its internal organization. Experiments performed during learning and on the trained network showed that: (i) the task could be learned by a three-layer sequential network; (ii) the network successfully generalized in trajectory space and adjusted the velocity profiles properly; (iii) the same task could not be learned by a linear network; (iv) after learning, the internal connections became organized into inhibitory and excitatory zones and encoded the main features of the training set; (v) the model was robust to noise on the input signals; (vi) the network exhibited attractor-dynamics properties; (vii) the network was able to solve the motorequivalence problem. A key feature of this work is the fact that the neural network was coupled to a mechanical model of a limb in which muscles are represented as springs. With this representation the model solved the problem of motor redundancy.     

Muscle activation and coactivation during five-time-sit-to-stand movement in patients undergoing total knee arthroplasty

Quadriceps weakness is prevalent with knee osteoarthritis (OA) and after total knee arthroplasty (TKA). To compensate for quadriceps dysfunction, patients often alter movement strategies. Little is known about muscle coordination during sit-to-stand (concentric) and stand-to-sit (eccentric) movements in the acute postoperative period. This investigation characterized the distribution of muscle activation between the concentric and eccentric phases during a five-time-sit-to-stand (FTSTS) movement in late stage OA and one month after TKA. Patients and healthy participants performed a FTSTS while recording bilateral ground reaction forces (GRFs) and electromyography (EMG). Concentric and eccentric ensemble averages of the GRF and EMG were calculated for the concentric and eccentric phases. Coactivation indices, integrated EMG, and GRF were calculated for each limb and phase. Patients demonstrated higher eccentric coactivation than the healthy group. Postoperative loading was higher in the nonsurgical limb. Postoperative quadriceps activity was lower in the concentric phase and higher in the eccentric phase than the healthy group. Higher coactivation in the patients resulted from sustained distribution of quadriceps activity throughout the eccentric phase. This indicated an inability to coordinate muscle firing when rapidly lowering to a chair and occurred despite unloading of the surgical limb. Although these patterns may serve as a protective strategy, they may also impede recovery of muscle function after TKA.     

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.     

Characteristics of locomotion regulation in man

It has been found by transforming experimental kinematic data to normal coordinates with calculating of muscle force moments during walking that the locomotor movements are regulated almost discontinuously at each degree of freedom of leg, so two piece constant parameters of control are switched few times during gait cycle. Therefore musculature acts like switched elastic constraints, and energy expenditure depends on the trajectories of movements essentially less than on the kinematic conditions displayed during fixed switchings.