Peculiarities of Activation of Muscles of the Shoulder Belt in Voluntary Two-Joint Movements of the Upper Limb

We studied coordination of central motor commands (СMCs) coming to muscles of the shoulder and shoulder belt in the course of single-joint and two-joint movements including flexion and extension of the elbow and shoulder joints. Characteristics of rectified and averaged EMGs recorded from a few muscles of the upper limb were considered correlates of the CMC parameters. Special attention was paid to coordination of CMCs coming to two-joint muscles that are able to function as common flexors (m. biceps brachii, caput breve, BBcb) and common extensors (m. triceps brachii, caput longum, TBcl) of the elbow and shoulder joints. Upper limb movements used in the tests included planar shifts of the arm from one spatial point to another resulting from either simultaneous changes in the angles of the shoulder and elbow joints or isolated sequential (two-stage) changes in these joint angles. As was found, shoulder muscles providing movements of the elbow with changes in the angle of the elbow joint, i.e., BBcb and TBcl, were also intensely involved in the performance of single-joint movements in the shoulder joint. The CMCs coming to two-joint muscles in the course of two-joint movements appeared, in the first approximation, as sums of the commands received by these muscles in the course of corresponding single-joint movements in the elbow and shoulder joints. Therefore, if we interpret the isolated forearm movement performed due to a change in the angle of the elbow joint as the main motor event, while the shoulder movement is considered the accessory one, we can conclude that realization of a two-joint movement of the upper-limb distal part is based on superposition of CMCs related to basic movements (main and accessory). Neirofiziologiya/Neurophysiology, Vol. 41, No. 1, pp. 48–56, January–February, 2009.     

Peculiarities of Activation of the Upper Limb Muscles in Realization of Two-Joint Movements in Humans

In tests on humans, we recorded EMG activity from the muscles flexing and extending the forearm and shoulder in the course of realization of sequential single-joint and simultaneous two-joint movements of the upper limb. As was shown, the shoulder muscles m. biceps brachii and m. triceps brachii are involved in flexion/extension of both elbow and shoulder joints. Central commands sent to the above muscles in the course of a two-joint movement could be considered a superposition of the central commands coming to the same muscles in realization of the corresponding sequential single-joint movements with the same changes in the angles of the elbow and shoulder joints. External loadings applied in the direction of extension of the elbow and shoulder joints induced, in general, similar changes in coordination of the activity of muscles moving the forearm and shoulder under conditions of both single-joint and two-joint movements. These facts allow us to suppose that coordination of the muscle activity in two-joint movements depends to a greater extent on the forces influencing limb links than on the mode of realization of the movements (two sequential single-joint movements vs a two-joint movement corresponding to the above motor events).     

Activation of the Shoulder Belt and Shoulder Muscles of Humans Related to Different Rates of Generation of Two-Joint Efforts by the Forearm

We studied coordination of central motor commands (CMCs) coming to the muscles that flex and extend the shoulder and elbow joints in the course of generation of voluntary isometric efforts of different directions by the forearm. Dependences of the characteristics of these commands on the direction of the effort and rate of its generation were analyzed. Amplitudes of rectified and averaged EMGs recorded from a number of shoulder belt and shoulder muscles were considered correlates of the CMC intensity. The development of the effort of a given direction and rate of rise was realized in the horizontal-plane operational space; the arm position corresponded to the 30 deg angle in the shoulder joint (external angle with respect to the frontal plane) and 90 deg angle in the elbow joint. We plotted sector diagrams of the relative changes in the level of dynamic and stationary phases of EMG activity of the studied muscles for the entire set of directions of the efforts generated with different rates of rise. In the course of formation of rapid two-joint isometric efforts, realization of nonsynergic motor tasks (extension of one joint and flexion of another one, and vice versa) required significant activation of muscles of different functional directions for both joints. Time organization of EMG activity of extensors and flexors of the shoulder and elbow joints related to the maximum and relatively rapid generation of the effort (rise time 0.12 to 0.13 and 0.25 sec, respectively) was rather complex and included dynamic and stationary phases. With these time parameters of generation of the efforts (both flexion and extension), the appearance at the stationary effort of 40 N was controlled based on coordinated interaction of dynamic phases of the activation of agonistic and antagonistic muscles. It is concluded that CMCs coming to extensors and flexors of both joints upon generation of rapid isometric efforts are rather similar in their parameters to those under conditions of realization of the forearm movements in the space in an isotonic mode.     

Central Activation of the Upper Limb Muscles in Humans Related to Creation of an Isometric Effort: Dependence on the Position of the Point of Force Application within the Operational Space

We examined the peculiarities of central coordination of motor commands coming to the muscles of the shoulder belt and shoulder in the course of generation of targeted isometric efforts by the arm. The dependence of these commands on changes in the effort direction and position of the forearm within the working space were analyzed. The intensity of the central commands was estimated according to the amplitudes of rectified and averaged EMGs recorded from the corresponding muscles. Sector diagrams of EMG activity of the above muscles depending on the direction of the effort vector, EV, were plotted [1]. Preferential sectors of activity where the efforts were formed due to activation of definite functional muscle groups were identified. As was found, the direction of these sectors depends significantly on the EV orientation. Differences between the patterns of coactivation of the examined muscles were demonstrated. Organization of the motor commands under conditions of creation of extensor efforts is distinguished by a more complex pattern than that related to flexor efforts. In the former case, the activity of extensor muscles is accompanied by more significant activation of the flexors.     

Peculiarities of Activation of the Shoulder Belt and Shoulder Muscles in Generation of Different-Direction Isometric Efforts by the Forearm

We studied central motor commands, CMCs, coming to the muscles that flex and extend the shoulder and elbow joints in the course of generation of voluntary isometric efforts of different directions by the forearm; the efforts were initiated according to a visual signal. Amplitudes of EMGs recorded from the muscles of the shoulder belt and shoulder and subjected to full-wave rectification and low-frequency filtration were considered correlates of the CMC intensity. An effort of the preset direction was developed within the operational space of the horizontal plane with angles 30 deg in the shoulder joint (external angle with respect to the frontal plane) and 90 deg in the elbow joint. We plotted sector diagrams of the logarithmic coefficient of the intensity increment of EMGs of the above muscles for the entire set of directions of generated efforts with a 15- or 20-deg step. Orientations of the maxima of EMG activity of the given muscles were rather close to the directions of the maxima of the force moments generated by these muscles. In most cases, a shift of the direction by one gradation with respect to the EMG maximum in the respective muscle resulted in a significant decrease in the level of EMG activity. It is shown that preferential activation of the muscles agonistic with respect to the examined direction of the generated effort was, as a rule, accompanied by coactivation of the antagonist muscles. When “two-joint” isometric efforts are formed, realization of the socalled synergic muscle tasks (where prevailing contractions of the muscles of the same functional direction for both joints coincide, i.e., flexion-flexion or extension-extension) is organized in a simpler manner. The programs of “nonsynergic” contractions (flexion of one joint and extension of another one, or vice versa) are more complex. In different subjects, considerably dissimilar patterns of EMG activity in muscles influencing these joints could be observed.     

Changes in limb dynamics during the practice of rapid arm movements

In our study we examined Bernstein's hypothesis that practice alters the motor coordination among the muscular and passive joint moments. In particular, we conducted dynamical analyses of a human multisegmental movement during the practice of a task involving the upper extremity. Seven male human volunteers performed maximal-speed, unrestrained vertical arm movements whose upward and downward trajectories between two target endpoints required the hand to round a barrier, resulting in complex shoulder, elbow, and wrist joint movements. These movements were recorded by high-speed ciné film, and myopotentials from selected upper-extremity muscles were recorded. The arm was modeled as interconnected rigid bodies, so that dynamical interactions among the upper arm, forearm, and hand could be calculated. With practice, subjects achieved significantly shorter movement times. As movement times decreased, all joint-moment components (except gravity) increased, and the moment-time and EMG profiles were changed significantly. Particularly during reversals in movement direction, the changes in moment-time and EMG profiles were consistent with Bernstein's hypothesis relating practice effects and intralimb coordination: with practice, motor coordination was altered so that individuals employed reactive phenomena in such a way as to use muscular moments to counterbalance passive-interactive moments created by segment movements.     

Velocity-Based Planning of Rapid Elbow Movements Expands the Control Scheme of the Equilibrium Point Hypothesis

According to the equilibrium point hypothesis of voluntary motor control, control action of muscles is not explicitly computed, but rather arises as a consequence of interaction between moving equilibrium position, current kinematics and stiffness of the joint. This approach is attractive as it obviates the need to explicitly specify the forces controlling limb movements. However, many debatable aspects of this hypothesis remain in the manner of specification of the equilibrium point trajectory and muscle activation (or its stiffness), which elicits a restoring force toward the planned equilibrium trajectory. In this study, we expanded the framework of this hypothesis by assuming that the control system uses the velocity measure as the origin of subordinate variables scaling descending commands. The velocity command is translated into muscle control inputs by second order pattern generators, which yield reciprocal command and coactivation commands, and create alternating activation of the antagonistic muscles during movement and coactivation in the post-movement phase, respectively. The velocity command is also integrated to give a position command specifying a moving equilibrium point. This model is purely kinematics-dependent, since the descending commands needed to modulate the visco-elasticity of muscles are implicitly given by simple parametric specifications of the velocity command alone. The simulated movements of fast elbow single-joint movements corresponded well with measured data performed over a wide range of movement distances, in terms of both muscle excitations and kinematics. Our proposal on a synthesis for the equilibrium point approach and velocity command, may offer some insights into the control scheme of the single-joint arm movements.     

Maximum isometric arm forces in the horizontal plane

In this paper, we measured the maximum isometric force at the hand in eight directions in the horizontal plane and at five positions in the workplace. These endpoint forces were the result of shoulder horizontal adduction/abduction and elbow flexion/extension torques. We found that the normalized maximum forces of all the six subjects deviated less than 15%, despite intra-subject differences in muscle strength of more than a factor of two. The maximum forces were found to systematically depend on the force direction and on the hand position in the workspace. The largest forces were found in a direction approximately along the line connecting shoulder joint and hand, and the smallest forces perpendicular to that line, thereby forming an elliptically shaped pattern. The elongation of the pattern was the largest for those hand positions having the more extended elbow joint. By using a lumped six-muscle model, with two mono-articular muscle pairs and one bi-articular pair, we were able to predict the observed force patterns. Here, we assumed that one of the muscles generates its maximum force and the others adjust their output to point the endpoint force in the required direction. We used a principal component analysis of the surface EMGs of simultaneously measured representatives of four of the six muscles. With the same model, we were then able to determine the principal directions of all the six muscle groups.     

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     

Movements of the Forearm and Shoulder Performed against the Gravitation Force: Target Positioning under Kinesthetic Control

In humans, we tested targeted movements of the forearm and shoulder performed in the vertical direction (in a parallel manner with respect to the sagittal plane). Movements were realized, first, with the possibility for visual control of the coincidence of the angle of the limb link axis vs the vertical and the target angle value (using an optic system and video recording), and, second, in the absence of the above control. Movements including flexion (i.e., movement against the gravitation force) – extension of the limb link with an individually selected convenient velocity were initiated and terminated according to the presentation of permissive sound signals; simultaneously, EMGs were recorded from a few muscles flexing and extending the elbow and shoulder joints. We analyzed systematic errors of target positioning of the forearm and shoulder in movements realized exclusively under kinesthetic control. In the case of isolated flexion of the forearm for a 90 deg target angle, such errors in all members of the examined group (n = 11) were positive. These errors were, on average, 8.1 ± 0.7 deg without loading and reached 11.2 ± 0.9 deg with introduction of a 10 to 30 N additional loading on the forearm. Isolated movements of the shoulder for a 70 deg target angle (performed without loading, with full extension of the forearm and its voluntary fixation) were accompanied by positive errors of 18.3 ± 1.1 deg, on average. Both the movements and positioning were performed due to changes in the levels of activity of the flexor muscles, with minimum involvement of the antagonists. The nonlinear properties of the receptor apparatus responsible for the formation of a kinesthetic estimate of the joint angle (first of all, of muscle spindles) are a fundamental reason for positive errors of target positioning of the limb links realized under kinesthetic control in the absence of the visual one.     

Resistance training enhances the stability of sensorimotor coordination

Strategies for the control of human movement are constrained by the neuroanatomical characteristics of the motor system. In particular, there is evidence that the capacity of muscles for producing force has a strong influence on the stability of coordination in certain movement tasks. In the present experiment, our aim was to determine whether physiological adaptations that cause relatively long-lasting changes in the ability of muscles to produce force can influence the stability of coordination in a systematic manner. We assessed the effects of resistance training on the performance of a difficult coordination task that required participants to synchronize or syncopate movements of their index finger with an auditory metronome. Our results revealed that training that increased isometric finger strength also enhanced the stability of movement coordination. These changes were accompanied by alterations in muscle recruitment patterns. In particular, the trained muscles were recruited in a more consistent fashion following the programme of resistance training. These results indicate that resistance training produces functional adaptations of the neuroanatomical constraints that underlie the control of voluntary movement.     

Electrical activity of muscles during disturbances in the trajectory of single-joint motion in the arm

Horizontal "extension-flexion" movement of human arm at the elbow joint was studied. Humans performed movements in accordance with the instruction to ignore sudden trajectory changes that arose by the arm during lengthening of different elastic rods with friction. It was shown by multi-dimension regression analysis that the electric activity of biceps and triceps muscles of the shoulder was correlated first of all with the work of load and then with loading force, muscle work and other mechanic and myographic characteristics. One could suppose that information on energy quantities was contained in afferentation signals of the nervous system to correct the program commands on the spinal level. The purpose of correction was to preserve constant quantity of energy expenditure, as well as accuracy and duration of movement.     

Intermuscular coordination in a sprint push-off.

This study was designed to investigate the patterns of intermuscular coordination during a sprinting event. In previous research it was found that despite the indeterminacy problem of movement control, movements like vertical jumping, speed skating and cycling are performed in a stereotyped manner. It was hypothesized that this might be due to constraints associated with the transformation of joint rotations into the desired translation. The objective of the present study was to determine the extent to which the intermuscular coordination patterns during other movements also are performed in a stereotyped manner and, if that is true, whether this can be understood on the basis of such constraints. Seven elite sprint runners were instructed to execute an explosive sprinting dash. Ground reaction forces and cinematographic data were recorded for the second stance phase of the sprint. Simultaneously, electromyographic activity of nine leg muscles was recorded telemetrically. Linked-segment modeling was used to obtain net joint moments and net joint powers. Different athletes appeared to perform the sprint in a stereotyped manner. The muscle coordination pattern is characterized by a proximo to distal sequence in timing of the monoarticular muscles. When compared to the sequential pattern found in jumping, the biarticular hamstrings and rectus femoris muscles behave differently; in the sprint a more pronounced reciprocal activity between these muscles exists. The resulting movement pattern is characterized by a sequence of upper leg extension and plantar flexion. The observed sequence in timing of muscle activation patterns is aimed at solving the problems associated with the earlier identified geometrical and anatomical constraint. However, the coordination pattern cannot be fully understood on the basis of these constraints. A specific constraint is identified with respect to the direction of the ground reaction force, which explains the pronounced reciprocal activity of the biarticular hamstring and rectus femoris muscles. The intermuscular coordination pattern in the sprint can be seen as a compromise between the specific requirement of the sprint and the advantageous effect of a proximo to distal sequence as found previously for jumping.     

Muscles within muscles: Coordination of 19 muscle segments within three shoulder muscles during isometric motor tasks.

The aim of the present study was to determine how the intra-muscular segments of three shoulder muscles were coordinated to produce isometric force impulses around the shoulder joint and how muscle segment coordination was influenced by changes in movement direction, mechanical line of action and moment arm (ma). Twenty male subjects (mean age 22 years; range 18-30 years) with no known history of shoulder pathologies, volunteered to participate in this experiment. Utilising an electromyographic technique, the timing and intensity of contraction within 19 muscle segments of three superficial shoulder muscles (Pectoralis Major, Deltoid and Latissimus Dorsi) were studied and compared during the production of rapid (e.g. approximately 400ms time to peak) isometric force impulses in four different movement directions of the shoulder joint (flexion, extension, abduction and adduction). The results of this investigation have suggested that the timing and intensity of each muscle segment's activation was coordinated across muscles and influenced by the muscle segment's moment arm and its mechanical line of action in relation to the intended direction of shoulder movement (e.g. flexion, extension, abduction or adduction). There was also evidence that motor unit task groups were formed for individual motor tasks which comprise motor units from both adjacent and distant muscles. It was also confirmed that for any particular motor task, individual muscle segments can be functionally classified as prime mover, synergist or antagonist - classifications which are flexible from one movement to the next.     

Mechanical energy expenditure on human movement and the anthropomorphic mechanism]

Mechanical energy expenditures of the man and anthropomorphic locomotion machine during movement are compared theoretically. Sources of the mechanical energy affecting movement of human's lower extremity are modelled by 8 muscles, 3 of which are the two-joint muscles. The model of the lower extremity of anthropomorphic locomotion machine is moved by joint moments. It was shown that in the same movement the model of the human lower extremity can spend less mechanical energy than that of the model of the anthropomorphic locomotion machine. It is caused by the presence of two-joint muscles in the first model. Such an economy of mechanical energy expenditures realized by the two-joint muscle is possible at simultaneous execution of three conditions: 1) signs of the muscle powers, which are produced by that muscle at both joints, are opposite; 2) moments produced by that muscle at each of both joints have the same direction with the joint moments at these joints; 3) one-joint antagonistic muscles are not active. An expression which makes it possible to estimate the mechanical energy savings by the two-joint muscles during humans' movement was developed.     

Adaptation to fatigue of long duration in human wrist movements.

Subjects made fast, accurate, consistent wrist flexions under normal conditions and under conditions of low-frequency fatigue. Movements made 1 h after fatiguing exercise were indistinguishable from those made before exercise, even though twitch tensions were only approximately 60% of their fresh values. Electromyograms (EMGs) recorded from the fatigued muscles were, however, different from those recorded before exercise. EMGs during unfatigued movements showed multiple bursts typical for rapid movements. In the presence of low-frequency fatigue, the duration of the first burst was longer than that under normal conditions, and its onset occurred earlier relative to the initiation of movement. The area of the second agonist burst and, in some cases, the antagonist burst, was increased, although changes in their timings were unclear. We conclude that subjects adapted to low-frequency fatigue by changing the neural patterns controlling their muscles and present a simple model of excitation-contraction coupling that demonstrates how the observed changes in excitation can produce the same kinematics.     

Effects of postural muscle fatigue on the relation between segmental posture and movement.

The purpose of this study was to examine whether fatigue of postural muscles might influence the coordination between segmental posture and movement. Seven healthy adults performed series of fifteen fast wrist flexions and extensions while being instructed to keep a dominant upper limb posture as constant as possible. These series of voluntary movements were performed before and after a fatiguing submaximal isometric elbow flexion, and also with or without the help of an elbow support. Surface EMG from muscles Delto?deus anterior, Biceps brachii, Triceps brachii, Flexor carpi ulnaris, Extensor carpi radialis were recorded simultaneously with wrist, elbow and shoulder accelerations and wrist and elbow displacements. Fatigue was evidenced by a shift of the elbow and shoulder muscles EMG spectra towards low frequencies. Kinematics of wrist movements and corresponding activations of wrist prime-movers, as well as the background of postural muscle activation before wrist movement were not modified. There were only slight changes in timing of postural muscle activations. These data indicate that postural fatigue induced by a low-level isometric contraction has no effect on voluntary movement and requires no dramatic adaptation in postural control.     

Mutual influences of upper and lower extrimities during cyclic movements

The possibility of muscle activation of passive arm during its cyclic movements, imposed by active movements of contralateral arm or by experimenter was studied, as well as the influence of lower extremities cyclic movements onto arm muscles activity. In addition to that the activity of legs muscles was estimated in dependence on motor task condition for arms. Ten healthy supine subjects carried out opposite movements of arms with and without stepping-like movements of both legs. The experiment included three conditions for arm movements: 1) the active movements of both arms; 2) the active movements of one arm, when other entirely passive arm participated in the movement by force; 3) passive arm movement caused by experimenter. In the condition 2) additional load on active arm was applied (30 N and 60 N). In all three conditions the experiment was carried out with arms movements only or together with legs movements. The capability of passive moving arm muscles activation depended on increasing afferent inflow from muscles of contralateral arm was demonstrated. Emerging electrical activity was modulated in the arms movements cycle and depended on the degree of active arm loading. During combined active movements of arms and legs the reduction of activity in the flexor muscles of shoulder and forearm was observed. Concomitant arms movements increased the magnitude ofelectromiographic bursts during passive stepping-like movements in the most of recorded muscles, and the same increasing was only observed in biceps femoris and tibialis anterior muscles during active legs movement. The increasing of loading of one arm caused essential augmentation of EMG-activity in the majority of recording legs muscles. The data obtained are the additional proof of existence of functionally significant neuronal interaction both between arms and between upper and lower extremities, which is evidently depend on the intraspinal neuronal connections.     

Torque and force production during shoulder external rotation: differences between transverse and sagittal planes

In joints with 3 degrees of freedom, such as the shoulder joint, the association of different movements results in changes in the behavior of the moment arm of the muscles. The capacity of torque production for the same movement can be changed when movements take place in a different plane. The objective of this study is to quantify differences between torque production and resultant force estimated during the shoulder external rotation in two movement planes: the transverse and sagittal planes (with 90 degrees of shoulder abduction). Eight individuals were evaluated using an isokinetic dynamometer and an eletrogoniometer for movements in the transverse plane and six individuals for movements in the sagittal plane. The results showed that the execution of the external rotation in the sagittal plane allowed greater torque magnitudes and resultant force compared with those in the transverse plane, probably owing to a prestretching of infraspinatus and teres minor.