Biomechanical imperatives in the neural control of locomotion (original) (raw)
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Integrative and Comparative Biology
Navigating complex terrains requires dynamic interactions between the substrate, musculoskeletal, and sensorimotor systems. Current perturbation studies have mostly used visible terrain height perturbations, which do not allow us to distinguish among the neuromechanical contributions of feedforward control, feedback-mediated, and mechanical perturbation responses. Here, we use treadmill-belt speed perturbations to induce a targeted perturbation to foot speed only, and without terrain-induced changes in joint posture and leg loading at stance onset. Based on previous studies suggesting a proximo-distal gradient in neuromechanical control, we hypothesized that distal joints would exhibit larger changes in joint kinematics, compared to proximal joints. Additionally, we expected birds to use feedforward strategies to increase the intrinsic stability of gait. To test these hypotheses, seven adult guinea fowl were video recorded while walking on a motorized treadmill, during both steady a...
Biomechanical and physiological aspects of legged locomotion in humans
European Journal of Applied Physiology, 2003
Walking and running, the two basic gaits used by man, are very complex movements. They can, however, be described using two simple models: an inverted pendulum and a spring. Muscles must contract at each step to move the body segments in the proper sequence but the work done is, in part, relieved by the interplay of mechanical energies, potential and kinetic in walking, and elastic in running. This explains why there is an optimal speed of walking (minimal metabolic cost of about 2 J.kg–1·m–1 at about 1.11 m.s–1) and why the cost of running is constant and independent of speed (about 4 J.kg–1.m–1). Historically, the mechanical work of locomotion has been divided into external and internal work. The former is the work done to raise and accelerate the body centre of mass (m) within the environment, the latter is the work done to accelerate the body segments with respect to the centre of m. The total work has been calculated, somewhat arbitrarily, as the sum of the two. While the changes of potential and kinetic energies can be accurately measured, the contribution of the elastic energy cannot easily be assessed, nor can the true work performed by the muscles. Many factors can affect the work of locomotion - the gradient of the terrain, body size (height and body m), and gravity. The partitioning of positive and negative work and their different efficiencies explain why the most economical gradient is about –10% (1.1 J.kg–1.m–1 at 1.3 m.s–1 for walking, and 3.1 J.kg–1.m–1 at between 3 and 4 m·s–1 for running). The mechanics of walking of children, pigmies and dwarfs, in particular the recovery of energy at each step, is not different from that of taller (normal sized) individuals when the speed is expressed in dynamically equivalent terms (Froude number). An extra load, external or internal (obesity) affects internal and external work according to the distribution of the added m. Different gravitational environments determine the optimal speed of walking and the speed of transition from walking to running: at more than 1 g it is easier to walk than to run, and it is the opposite at less than 1 g. Passive aids, such as skis or skates, allow an increase in the speed of progression, but the mechanics of the locomotion cannot be simply described using the models for walking and running because step frequency, the proportion of step duration during which the foot is in contact with the ground, the position of the limbs, the force exerted on the ground and the time of its application are all different.
Acta Physiologica Scandinavica, 1985
Knowledge of adaptations to changes in speed and mode of progression (walking-running) in human locomotion is important for an understanding of underlying neural control mechanisms and allows a comparison with more detailed animal studies. Leg movements and muscle activity patterns were studied in ten healthy males (19–29 yr) during level walking (0.4-3.0 m-s-1) and running (1.0–9.0 m-s-1) on a motor-driven treadmill. Movements were recorded in the sagittal plane with a Selspot optoelectronic system. Recordings of EMG were made from seven different muscles of one leg by means of surface electrodes. Durations, amplitudes and relative phase relationships of angular displacements and EMG activity were analysed in relation to different phases of the stride cycle (defined by the leg movements). The durations of the entire stride cycle and of the support phase were found to decrease curvilinearly with velocity. Swing and support phase durations were linearly related to cycle duration in walking, and curvilinearly related in running. The characteristic occurrence of double support phases in walking was also seen in very slow running. Support length increased with speed up to about i,2 m both in walking and running, but was longer in walking at the same velocity. Increases in net angular displacements were largest for hip movements and for knee flexion-extension during the swing phase in running. With increasing velocity a clear shift in relative rectus femoris activity occurred from knee extension to hip flexion. Gastrocnemius lateralis (LG) was co-activated with the other leg extensors prior to foot contact in running, whereas in walking LG was not turned on until later in the support phase. The ankle flexor tibialis anterior had its main peak of activity after touch-down in walking and before touch-down in running. The same basic structure of the stride cycle as in other animals suggests similarities in the underlying neural control. Human speed adaptation is distinguished primarily by an increase in both frequency and amplitude of leg movements and by a possibility of changing between a walking and a running type of movement pattern.