We studied the changes of vertical contact forces, lower limb kinematics, and electromyographic activity (EMG) at different speeds and gravitational loads. To this end healthy subjects were asked to walk on a motorized treadmill while the percentage of body weight unloaded (body weight support, BWS) was modified in steps by means of a well-characterized unloading system. BWS was set at 0, 35, 50, 75, 95, or 100% of body weight. Walking speed was 0.7, 1.1, 2, 3, or 5 km/h. We found that changing BWS between 0 and 95% resulted in drastic changes of kinetic parameters but in limited changes of the kinematic coordination. In particular, the peak vertical contact forces decreased proportionally to BWS; at 95%-BWS they were 20-fold smaller than at 0% and were applied at the forefoot only. Also, there were considerable changes of the amplitude of EMG activity of all tested lower limb muscles and a complex re-organization of the pattern of activity of thigh muscles. By contrast, the corresponding variation of the parameters that describe shape and variability of the foot path was very limited, always <30% of the corresponding values at 0 BWS. Moreover, the planar co-variation of the elevation angles was obeyed at all speed and BWS values. Minimum variance of limb trajectory occurred at 3 km/h. At 100% BWS, subjects stepped in the air, their feet oscillating back and forth just above but never contacting the treadmill. In this case, step-to-step variability of foot path was much greater than at all other BWS levels but was restored to lower values when minimal surrogate contact forces were provided during the "stance" phase. The results did not depend on the specific instruction given to the subject. Therefore we conclude that minimal contact forces are sufficient for accurate foot trajectory control.

We studied the effect of vibratory stimulation of different leg muscles [bilateral quadriceps (Q), hamstring (HS) muscles, triceps surae (TS), and tibialis anterior (TA)] in seven normal subjects during 1) quiet standing, 2) stepping in place movements, and 3) walking on the treadmill. The experiments were performed in a dimly illuminated room, and the subjects were given the instruction not to resist the applied perturbation. In one condition the velocity of the treadmill was controlled by a feedback from the subject's current position. In normal standing, TA vibration elicited a prominent forward body tilt, whereas HS and TS vibration elicited backward trunk or whole body inclination, respectively. Q vibration had little effect. During stepping in place, continuous HS vibration produced an involuntary forward stepping at about 0.3 m s(-1) without modifying the stepping frequency. When the subjects (with eyes closed) kept a hand contact with an external still object, they did not move forward but perceived an illusory forward leg flexion relative to the trunk. Q, TS, and TA vibration did not cause any systematic body translation nor illusory changes in body configuration. In treadmill locomotion, HS vibration produced an involuntary steplike increase of walking speed (by 0.1-0.6 m.s(-1)). Continuous vibration elicited larger speed increments than phasic stimulation during swing or stance phase. For phasic stimulation, HS vibration tended to be more effective when applied during swing than during stance phase. Q, TA, and TS vibration had little if any effect. Vibration of thigh muscles altered the walking speed depending on the direction of progression. During backward locomotion, the walking speed tended to decrease after HS vibration, whereas it significantly increased after Q vibration. Thus the influence of leg muscle vibration on stepping in place and locomotion differed significantly from that on normal posture. We suggest that the proprioceptive input from thigh muscles may convey information about the velocity of the foot movement relative to the trunk.

Mature locomotion involves modular spinal drives generating a set of fundamental patterns of motoneuron activation, each timed at a specific phase of locomotor cycles and associated with a stable muscle synergy. How locomotor modules develop and to what extent they depend on prior experience or intrinsic programs remains unclear. To address these issues, we herein leverage the presence at birth of two types of locomotor-like movements, spontaneous kicking and weight-bearing stepping. The former is expressed thousands of times in utero and postnatally, whereas the latter is elicited de novo by placing the newborn on the ground for the first time. We found that the neuromuscular modules of stepping and kicking differ substantially. Neonates kicked with an adult-like number of temporal activation patterns, which lacked a stable association with systematic muscle synergies across movements. However, on the ground neonates stepped with fewer temporal patterns but all structured in stable synergies. Since kicking and ground-stepping coexist at birth, switching between the two behaviors may depend on a dynamic reconfiguration of the underlying neural circuits as a function of sensory feedback from surface contact. We tracked the development of ground-stepping in 4- to 48-mo-old infants and found that, after the age of 6 mo, the number of temporal patterns increased progressively, reaching adult-like conformation only after independent walking was established. We surmise that mature locomotor modules may derive by combining the multiple patterns of repeated kicking, on the one hand, with synergies resulting from fractionation of those revealed by sporadic weight-bearing stepping, on the other hand.

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When a toddler starts to walk without support, gait kinematics and electromyographic (EMG) activity differ from those of older children and the body displays considerable oscillations due to poor equilibrium. Postural instability clearly affects motor patterns in adults, but does instability explain why toddlers walk with a different gait? Here we addressed this question by comparing kinematics and EMGs in toddlers performing their first independent steps with or without hand or trunk support. Hand support significantly improved postural stability and some general gait parameters, reducing percent of falls, step width, lateral hip deviations and trunk oscillations. However, the kinematic and EMG patterns were unaffected by increased postural stability. In particular, the co-variance of the angular motion of the lower limb segments, the pattern of bilateral coordination of the vertical movement of the two hip joints, high variability of the foot path, the elliptic or single peak trajectory of the foot in the swing phase, and characteristic EMG bursts at foot contact remained idiosyncratic of toddler locomotion. Instead the toddler pattern shared fundamental features with adult stepping in place, suggesting that toddlers implement a mixed locomotor strategy, combining forward progression with elements of stepping in place. Furthermore, gait kinematics remained basically unchanged until the occurrence of the first unsupported steps and rapidly matured thereafter. We conclude that idiosyncratic features in newly walking toddlers do not simply result from undeveloped balance control but may represent an innate kinematic template of stepping.