Automatic control of limb movement and posture

Studies are reviewed that address the problem of the variables controlled by the central nervous system in the maintenance of body posture and limb movement against disturbing forces. The role of global variables of control, which take into account the dynamic state of the limb, is discussed. Neural substrates that are involved in the distributed control of kinematic and dynamic parameters are also considered.     

Closed-loop and open-loop control of posture and movement during human trunk bending

Closed-loop (CL) and open-loop (OL) types of motor control during human forward upper trunk bending are investigated. A two-joint (hip and ankle) biomechanical model of the human body is used. The analysis is performed in terms of the movements along eigenvectors of the motion equation (“eigenmovements” or “natural synergies”). Two analyzed natural synergies are called “H-synergy” (Hip) and “A-synergy” (Ankle) according to the dominant joint in each of these synergies. Parameters of CL control were estimated using a sudden support platform displacement applied during the movement execution. The CL gain in the H-synergy increased and in the A-synergy decreased during the movement as compared with the quiet standing. The analysis of the time course of OL control signal suggests that the H-synergy (responsible for the prime movement, i.e. bending per se) is controlled according to the EP theory whereas for the associated A-synergy (responsible for posture adjustment, i.e. equilibrium maintenance) muscle forces and gravity forces are balanced for any its final amplitude and therefore the EP theory is not applicable to its control.     

Simulation studies on the control of posture and movement in a multi-jointed limb

Simulation studies were performed to evaluate the effectiveness of different control schemes in stabilizing a multi-jointed limb (human arm) in response to force perturbations. The mechanical properties of the arm were modeled as a linear visco-elastic system and the effectiveness of negative feedback of angular position and torque was evaluated. The effectiveness of a given amount of position feedback depended strongly on the initial position of the arm and on the perturbation, while torque feedback was much more consistently effective in damping the motion of the limb.     

The influence of initial posture on the sit-to-stand movement

Head movements, ground reaction forces and electromyographic activity of selected muscles were recorded simultaneously from two subjects as they performed the sit-to-stand manouevre under a variety of conditions. The influence of initial leg posture on the magnitude of the various parameters under investigation was examined first. A preferred initial leg posture resulted in smaller magnitudes of head movement and ground reaction forces. EMG activity in some muscles, trapezius and erector spinae, decreased, while in others, quadriceps and hamstrings, it increased in the preferred leg posture. The decreases seen correlate with reductions in head movement observed. The effect of inhibiting habitual postural adjustments of the head and neck, by comparing "free" and "guided" movements was also examined. In guided movements there are significant reductions in head movement, ground reaction forces and EMG activity in trapezius, sternomastoid and erector spinae. It would appear that both initial leg posture and the abolition of habitual postural adjustment have a profound influence on the efficiency of the sit-to-stand manouevre. This preliminary study high-lights the practical importance of head posture in the diagnosis and treatment of movement disorders, as well as in movement education.     

Neuromechanics of muscle synergies for posture and movement

Recent research suggests that the nervous system controls muscles by activating flexible combinations of muscle synergies to produce a wide repertoire of movements. Muscle synergies are like building blocks, defining characteristic patterns of activation across multiple muscles that may be unique to each individual, but perform similar functions. The identification of muscle synergies has strong implications for the organization and structure of the nervous system, providing a mechanism by which task-level motor intentions are translated into detailed, low-level muscle activation patterns. Understanding the complex interplay between neural circuits and biomechanics that give rise to muscle synergies will be crucial to advancing our understanding of neural control mechanisms for movement.     

Effect of gravity and posture on lung mechanics.

The volume-pressure relationship of the lung was studied in six subjects on changing the gravity vector during parabolic flights and body posture. Lung recoil pressure decreased by approximately 2.7 cmH(2)O going from 1 to 0 vertical acceleration (G(z)), whereas it increased by approximately 3.5 cmH(2)O in 30 degrees tilted head-up and supine postures. No substantial change was found going from 1 to 1.8 G(z). Matching the changes in volume-pressure relationships of the lung and chest wall (previous data), results in a decrease in functional respiratory capacity of approximately 580 ml at 0 G(z) relative to 1 G(z) and of approximately 1,200 ml going to supine posture. Microgravity causes a decrease in lung and chest wall recoil pressures as it removes most of the distortion of lung parenchyma and thorax induced by changing gravity field and/or posture. Hypergravity does not greatly affect respiratory mechanics, suggesting that mechanical distortion is close to maximum already at 1 G(z). The end-expiratory volume during quiet breathing corresponds to the mechanical functional residual capacity in each condition.     

Interaction between the vertical posture and movement control systems during a quick voluntary arm raise

The study was aimed at a deeper understanding of the interaction between the system of vertical posture control and the system of voluntary movement control based on the analysis of postural muscle activity components resulting from the action of the former or the latter system. For this purpose, a quick arm raise was performed in the standing and sitting positions with body fixation at different levels, when the task of maintaining a vertical posture was simplified or completely eliminated. Under these conditions, the muscle activity associated with posture control was supposed to change, while the activity of muscles raising the arm was supposed to remain invariable. The results showed that the simplification of the posture control resulted in a decrease or elimination of anticipatory changes in the activity of some muscles. However, most of the muscle activity variations were retained even in the sitting position, and these variations appeared simultaneously with the activity of muscles raising the arm. The so-called “anticipatory postural activity” during an arm raise in a normal standing position is supposed to consist of two components: an initial component reflecting the work of the posture control system and a later component reflecting the work of the movement control system. It is suggested that the planning of muscle activity and exchange of information between these two systems take place only before the beginning of the movement; after that, they act independently and in parallel.     

Steady-state force-velocity relation in human multi-joint movement determined with force clamp analysis

To study the force-velocity characteristics of human knee-hip extension movement, a dynamometer, in which force was controlled by a servo system, was developed. Seated subjects pressed either bilaterally or unilaterally a force plate, a horizontal position of which was servo-controlled so as to equalize the measured force and a force command generated by a computer at a time resolution of 2 ms (force clamp). The force command was based on the relation between maximum isometric force and foot position within the range between 70% and 90% of "leg length" (LL: longitudinal distance between the sole of the foot and the hip joint), so that the same force relative to the maximum isometric force was consistently applied regardless of the foot position. By regulating the force according to this function, the force-velocity relation was determined. The force-velocity relation obtained was described by a linear function (n=17, r=-0.986 for 80% LL, r=-0.968 for 85% LL) within a range of force between 0.1 and 0.8F(0) (maximum isometric force). The maximum force extrapolated from the linear regression (F(max)) coincided with F(0) (n=17, F(0)/F(max)=1.00+/-0.09 for 80% LL and 1.00+/-0.20 for 85% LL). Also, the velocity at zero force (V(max)) was obtained from the extrapolation. When compared to the bilateral movements, unilateral movements gave rise to a smaller F(max) but the same V(max), suggesting that V(max) is independent of force and therefore represents the proper unloaded velocity. It is suggested that some neural mechanisms may be involved in the force-velocity relation of the knee-hip extension movement, and make it exhibit a linear appearance rather than a hyperbola.     

The cortical control of movement revisited

Recently, we found that electrical stimulation of motor cortex caused monkeys to make coordinated, complex movements. These evoked movements were arranged across the cortex in a map of spatial locations to which the hand moved. We suggest that some of the subdivisions previously described within primary motor and premotor cortex may represent different types of actions that monkeys tend to make in different regions of space. According to this view, primary and premotor cortex may fit together into a larger map of manual space.     

Load sensing and control of posture and locomotion

This article reviews recent findings on how forces are detected by sense organs of insect legs and how this information is integrated in control of posture and walking. These experiments have focused upon campaniform sensilla, receptors that detect forces as strains in the exoskeleton, and include studies of sensory discharges in freely moving animals and intracellular characterization of connectivity of afferent inputs in the central nervous system. These findings provide insights into how campaniform sensilla can contribute to the adjustment of motor outputs to changes in load. In this review we discuss (1) anatomy of the receptors and their activities in freely moving insects, (2) mechanisms by which inputs are incorporated into motor outputs and (3) the integration of sensory signals of diverse modalities. The discharges of some groups of receptors can encode body load when standing. Responses are also correlated with muscle-generated forces during specific times in walking. These activities can enhance motor outputs through reflexes and can affect the timing of motoneuron firing through inputs to pattern generating interneurons. Flexibility in the system is also provided by interactions of afferent inputs at several levels. These mechanisms can contribute to the adaptability of insect locomotion to diverse terrains and environments.     

Optimal coordination and control of posture and movements

This paper presents a theoretical model of stability and coordination of posture and locomotion, together with algorithms for continuous-time quadratic optimization of motion control. Explicit solutions to the Hamilton–Jacobi equation for optimal control of rigid-body motion are obtained by solving an algebraic matrix equation. The stability is investigated with Lyapunov function theory and it is shown that global asymptotic stability holds. It is also shown how optimal control and adaptive control may act in concert in the case of unknown or uncertain system parameters. The solution describes motion strategies of minimum effort and variance. The proposed optimal control is formulated to be suitable as a posture and movement model for experimental validation and verification. The combination of adaptive and optimal control makes this algorithm a candidate for coordination and control of functional neuromuscular stimulation as well as of prostheses. Validation examples with experimental data are provided.     

Effects of posture on the respiratory mechanics of the chick embryo

In the chicken embryo, pulmonary ventilation and pulmonary gas exchange begin approximately one day before the completion of hatching. We asked to what extent the posture inside the egg, and the presence of the eggshell and membranes, may alter the mechanical behaviour of the respiratory system. The passive mechanical properties of the respiratory system were studied in chicken embryos during the internal pipping phase (rupture of the air cell) or the external pipping phase (hole in the eggshell). Tracheal pressure and changes in lung volume were recorded during mechanical ventilation, first, with the embryo curled up inside the egg, then again after exteriorization from the eggshell. In the internal pippers, respiratory system compliance increased and expiratory resistance decreased after exteriorization, whereas the mean inspiratory impedance did not change. In the external pippers, exteriorization had no significant effects on respiratory compliance, resistance, or impedance, and the values were similar to those of newly hatched chicks. We conclude that, in the chicken embryo, at a time when pulmonary ventilation becomes an important mechanism for gas exchange, the curled up posture inside the egg does not provide any significant mechanical constraint to breathing.     

Characteristics of the pressure center movement under upright posture regulation

The hypothesis was put forward that, along with the regulation of mass center projection, the system of upright posture control stabilizes the deviation of pressure center from the position of the mass center projection. The regularities in the behavior of the trajectories of pressure center and mass center projection were analysed. Experimental evidence was obtained supporting the validity of the hypothesis. The structure of the control system that corresponds to the new understanding of the variables being regulated during the maintenance of vertical posture was considered.     

Mechanisms of reference posture correction in the system of upright posture control

It was earlier shown that ultraslow tilts of the support under quiet standing conditions evoke an unusual response reflecting the operation of compensatory mechanisms: postural sway is a superposition of postural oscillations typical of quiet standing and greater, slower inclinations of the body caused by the tilt. This may be explained by the presence of two hierarchical levels of upright posture control: real-time control compensates for small deviations of the body from the reference posture prescribed by presetting control. Mathematical simulation methods have been used to study the mechanisms of reference posture control. The results are compared with available experimental data. It is demonstrated that the reference posture can be corrected according to the gravitational vertical with the use of a kinesthetic reference alone. It is hypothesized that, when correcting the reference posture, the nervous system “assumes” the support to be immobile. The afferent input from sole pressure receptors is an important factor in reference posture correction. The advantages of the putative two-level control over control based on an explicit internal model are discussed.     

The supplementary motor area is implicated in the coordination between posture and movement in man

The aim of the present experimental series was to investigate the central organization of the coordination between posture and movement in a bimanual load lifting task. The seated subject was instructed to maintain horizontal one forearm (postural arm) which was loaded with a 1 kg weight. The unloading was performed either by the experimenter (imposed unloading) or by a voluntary movement of the other arm (bimanual unloading). With the bimanual unloading, the movement control was accompanied by an anticipatory adjustment of the postural forearm flexors activity, which resulted in the maintenance of the forearm position despite the unloading. No change in the anticipatory postural adjustment was observed in one patient with complete callosal section. It was reduced in 5 patients with lesion of the SMA region, but only when the postural forearm was contralateral to the lesion. It is suggested that the SMA region contralateral to the postural forearm may select the circuits responsible for the phasic postural adjustments which are necessary to ensure postural maintenance, whereas the motor cortex contralateral to the voluntary movement controls both the movement and, via collaterals, the preselected circuits responsible for the associated postural adjustment.