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The neural locomotor system strongly relies on spinal circuitries. Yet, the control of bipedal gait is accompanied by activity in motor cortex. In human gait control, the functional interaction between these cortical contributions and their spinal counterparts are largely elusive. We focused on four spinal activation patterns during walking and explored their cortical signatures in toddlers and adults. In both groups, cortico-spinal coherence analysis revealed activity in primary motor cortex to be closely related to two of the four spinal patterns. Their corresponding muscle synergies are known to develop around the onset of independent walking. By hypothesis, the cortex hence contributes to the emergence of these synergies. In contrast, the other two spinal patterns investigated here resembled those present during newborn stepping. As expected, they did not show any cortical involvement. Together, our findings suggest a crucial role of motor cortex for independent walking in humans.

New-borns can step when supported for about 70–80% of their own body weight. Gravity-related sensorimotor information might be an important factor in developing the ability to walk independently. We explored how body weight support alters motor control in toddlers during the first independent steps and in toddlers with about half a year of walking experience. Sixteen different typically developing children were assessed during (un)supported walking on a running treadmill. Electromyography of 18–24 bilateral leg and back muscles and vertical ground reaction forces were recorded. Strides were grouped into four levels of body weight support ranging from no (

The aim of this study was to explore the underlying age-related differences in dynamic motor control during different step ascent conditions using muscle synergy analysis. Eleven older women (67.0 y &plusmn, 2.5) and ten young women (22.5 y &plusmn, 1.6) performed stepping in forward and lateral directions at step heights of 10, 20 and 30 cm. Surface electromyography was obtained from 10 lower limb and torso muscles. Non-negative matrix factorization was used to identify sets of (n) synergies across age groups and stepping conditions. In addition, variance accounted for (VAF) by the detected number of synergies was compared to assess complexity of motor control. Finally, correlation coefficients of muscle weightings and between-subject variability of the temporal activation patterns were calculated and compared between age groups and stepping conditions. Four synergies accounted for &gt, 85% VAF across age groups and stepping conditions. Age and step height showed a significant negative correlation with VAF during forward stepping but not lateral stepping, with lower VAF indicating higher synergy complexity. Muscle weightings showed higher similarity across step heights in older compared to young women. Neuromuscular control of young and community-dwelling older women could not be differentiated based on the number of synergies extracted. Additional analyses of synergy structure and complexity revealed subtle age- and step-height-related differences, indicating that older women rely on more complex neuromuscular control strategies.

The ability to walk without support usually develops in the first year of a typically developing toddler’s life and matures further in the following years. Mature walking is characterized by the correct timing of the different gait phases that make up a full gait cycle formed by stance, swing, and double support phases. The harmony of mature walking is given by a specific self-similar structure of the ratios between the durations of these phases (stride/stance, stance/swing, swing/double support), that in adults all converge to the golden ratio (phi, about 1.618). The aim of this longitudinal, prospective, experimental study was to investigate the evolution of this gait harmonic structure in the transition from supported to independent walking. In total, 27 children were recruited and recorded at various stages of locomotor development, ranging from neonatal stepping up to an independent walking experience of about six months. Differently from walking speed that progressively increased with age, the gait phase ratios started to converge to golden value only after the first independent steps, suggesting a relation to the maturation of the walking experience. The independent walking experience seems to represent a trigger for the evolution of a locomotor harmonic pattern in toddlers learning to walk.

Peripheral vision is often considered vital in (combat) sports, yet most experimental paradigms (e.g., eye tracking) ignore peripheral information or struggle to make inferences about the role of peripheral vision in an in-situ performance environment. This study aimed to determine where visual information is located in the peripheral field during an in-situ combat sports task. Eight advanced judokas competed in grip-fighting exchanges while wearing a mobile eye-tracker to locate gaze direction. Three-dimensional position data of the head and hands were tracked using a VICON motion capture system. Gaze analysis through automatic feature detection showed that participants predominantly fixated on their opponent’s chest. Kinematic data were used to calculate the angles between the opponent’s hands and the gaze-anchor point on the chest of the opponent. Results revealed a nonlinear relationship between visual field (VF) size and visibility of the hands, with athletes needing a VF of at least 30–40 degrees radius to simultaneously monitor both hands of the opponent most of the time. These findings hold implications for the regulation of Paralympic judo for athletes with vision impairment, suggesting that a less severe degree of impairment should be required to qualify than the current criterion of 20 degrees radius.

AIM: To determine if muscle synergy structure (activations and weights) differs between gait patterns in children with spastic cerebral palsy (CP). METHOD: In this cross-sectional study, we classified 188 children with unilateral (n=82) or bilateral (n=106) spastic CP (mean age: 9y 5mo, SD: 4y 3mo, range: 3y 9mo-17y 7mo; 75 females; Gross Motor Function Classification System [GMFCS] level I: 106, GMFCS level II: 55, GMFCS level III: 27) into a minor deviations (n=34), drop foot (n=16), genu recurvatum (n=26), apparent equinus (n=53), crouch (n=39), and jump gait pattern (n=20). Surface electromyography recordings from eight lower limb muscles of the most affected side were used to calculate synergies with weighted non-negative matrix factorization. We compared synergy activations and weights between the patterns. RESULTS: Synergy structure was similar between gait patterns, although weights differed in the more impaired children (crouch and jump gait) when compared to the other patterns. Variability in synergy structure between participants was high. INTERPRETATION: The similarity in synergy structure between gait patterns suggests a generic motor control strategy to compensate for the brain lesion. However, the differences in weights and high variability between participants indicate that this generic motor control strategy might be individualized and dependent on impairment level. ispartof: DEVELOPMENTAL MEDICINE AND CHILD NEUROLOGY vol:64 issue:4 pages:462-468 ispartof: location:England status: published

Muscle synergies reflect the presence of a common neural input to multiple muscles. Steering small sets of synergies is commonly believed to simplify the control of complex motor tasks like walking and running. When these locomotor patterns emerge, it is likely that synergies emerge as well. We hence hypothesized that in children learning to run the number of accompanying synergies increases and that some of the synergies’ activities display a temporal shift related to a reduced stance phase as observed in adults. We investigated the development of locomotion in 23 children aged 2–9 years of age and compared them with seven young adults. Muscle activity of 15 bilateral leg, trunk, and arm muscles, ground reaction forces, and kinematics were recorded during comfortable treadmill walking and running, followed by a muscle synergy analysis. We found that toddlers (2–3.5 years) and preschoolers (3.5–6.5 years) utilize a “walk-run strategy” when learning to run: they managed the fastest speeds on the treadmill by combining double support (DS) and flight phases (FPs). In particular the activity duration of the medial gastrocnemius muscle was weakly correlated with age. The number of synergies across groups and conditions needed to cover sufficient data variation ranged between four and eight. The number of synergies tended to be smaller in toddlers than it did in preschoolers and school-age children but the adults had the lowest number for both conditions. Against our expectations, the age groups did not differ significantly in the timing or duration of synergies. We believe that the increase in the number of muscle synergies in older children relates to motor learning and exploration. The ability to run with a FP is clearly associated with an increase in the number of muscle synergies.

Early brain lesions which produce cerebral palsy (CP) may affect the development of walking. It is unclear whether or how neuromuscular control, as evaluated by muscle synergy analysis, differs in young children with CP compared to typically developing (TD) children with the same walking ability, before and after the onset of independent walking. Here we grouped twenty children with (high risk of) CP and twenty TD children (age 6.5–52.4 months) based on their walking ability, supported or independent walking. Muscle synergies were extracted from electromyography data of bilateral leg muscles using non-negative matrix factorization. Number, synergies’ structure and variability accounted for when extracting one (VAF1 ) or two (VAF2 ) synergies were compared between CP and TD. Children in the CP group recruited fewer synergies with higher VAF1 and VAF2 compared to TD children in the supported and independent walking group. The most affected side in children with asymmetric CP walking independently recruited fewer synergies with higher VAF1 compared to the least affected side. Our findings suggest that early brain lesions result in early alterations of neuromuscular control, specific for the most affected side in asymmetric CP.

In this chapter, we explore the use of motion tracking methodology in developmental research. With motion tracking, also called motion capture, human movements can be precisely recorded and analyzed. Motion tracking provides developmental researchers with objective measurements of motor and (socio-)cognitive development. It can further be used to create carefully-controlled stimuli videos and can offer means of measuring development outside of the lab. We discuss three types of motion tracking that lend themselves to developmental applications. First, marker-based systems track optical or electromagnetic markers or sensors placed on the body and offer high accuracy measurements. Second, markerless methods entail image processing of videos to track the movement of bodies without participants being hindered by physical markers. Third, inertial motion tracking measures three-dimensional movements and can be used in a variety of settings. The chapter concludes by examining three example topics from developmental literature in which motion tracking applications have contributed to our understanding of human development.

Purpose We sought to identify the developing maturity of walking and running in young children. We assessed gait patterns for the presence of flight and double support phases complemented by mechanical energetics. The corresponding classification outcomes were contrasted via a shotgun approach involving several potentially informative gait characteristics. A subsequent clustering turned out very effective to classify the degree of gait maturity. Methods Participants (22 typically developing children aged 2–9 years and 7 young, healthy adults) walked/ran on a treadmill at comfortable speeds. We determined double support and flight phases and the relationship between potential and kinetic energy oscillations of the center-of-mass. Based on the literature, we further incorporated a total of 93 gait characteristics (including the above-mentioned ones) and employed multivariate statistics comprising principal component analysis for data compression and hierarchical clustering for classification. Results While the ability to run including a flight phase increased with age, the flight phase did not reach 20% of the gait cycle. It seems that children use a walk-run-strategy when learning to run. Yet, the correlation strength between potential and kinetic energies saturated and so did the amount of recovered mechanical energy. Clustering the set of gait characteristics allowed for classifying gait in more detail. This defines a metric for maturity in terms of deviations from adult gait, which disagrees with chronological age. Conclusions The degree of gait maturity estimated statistically using various gait characteristics does not always relate directly to the chronological age of the child.

When walking speed is increased, the frequency ratio between the arm and leg swing switches spontaneously from 2:1 to 1:1. We examined whether these switches are accompanied by changes in functional connectivity between multiple muscles. Subjects walked on a treadmill with their arms swinging along their body while kinematics and surface electromyography (EMG) of 26 bilateral muscles across the body were recorded. Walking speed was varied from very slow to normal. We decomposed EMG envelopes and intermuscular coherence spectra using non-negative matrix factorization (NMF), and the resulting modes were combined into multiplex networks and analyzed for their community structure. We found five relevant muscle synergies that significantly differed in activation patterns between 1:1 and 2:1 arm-leg coordination and the transition period between them. The corresponding multiplex network contained a single module indicating pronounced muscle co-activation patterns across the whole body during a gait cycle. NMF of the coherence spectra distinguished three EMG frequency bands: 4-8, 8-22, and 22-60 Hz. The community structure of the multiplex network revealed four modules, which clustered functional and anatomical linked muscles across modes of coordination. Intermuscular coherence at 4-22 Hz between upper and lower body and within the legs was particularly pronounced for 1:1 arm-leg coordination and was diminished when switching between modes of coordination. These findings suggest that the stability of arm-leg coordination is associated with modulations in long-distant neuromuscular connectivity.

Contrary to cats and primates, cortical contribution to hindlimb locomotor movements is not critical in rats. However, the importance of the motor cortex to regain locomotion after neurological disorders in rats suggests that cortical engagement in hindlimb motor control may depend on the behavioral context. To investigate this possibility, we recorded whole-body kinematics, muscle synergies, and hindlimb motor cortex modulation in freely moving rats performing a range of natural locomotor procedures. We found that the activation of hindlimb motor cortex preceded gait initiation. During overground locomotion, the motor cortex exhibited consistent neuronal population responses that were synchronized with the spatiotemporal activation of hindlimb motoneurons. Behaviors requiring enhanced muscle activity or skilled paw placement correlated with substantial adjustment in neuronal population responses. In contrast, all rats exhibited a reduction of cortical activity during more automated behavior, such as stepping on a treadmill. Despite the facultative role of the motor cortex in the production of locomotion in rats, these results show that the encoding of hindlimb features in motor cortex dynamics is comparable in rats and cats. However, the extent of motor cortex modulations appears linked to the degree of volitional engagement and complexity of the task, reemphasizing the importance of goal-directed behaviors for motor control studies, rehabilitation, and neuroprosthetics.SIGNIFICANCE STATEMENTWe mapped the neuronal population responses in the hindlimb motor cortex to hindlimb kinematics and hindlimb muscle synergies across a spectrum of natural locomotion behaviors. Robust task-specific neuronal population responses revealed that the rat motor cortex displays similar modulation as other mammals during locomotion. However, the reduced motor cortex activity during more automated behaviors suggests a relationship between the degree of engagement and task complexity. This relationship emphasizes the importance of the behavioral procedure to engage the motor cortex during motor control studies, gait rehabilitation, and locomotor neuroprosthetic developments in rats.

Vision can improve bipedal upright stability during standing and locomotion. However, during locomotion, vision supports additional behaviors such as gait cycle modulation, navigation, and obstacle avoidance. Here, we investigate how the multiple roles of vision are reflected in the dynamics of trunk control as the neural control problem changes from a fixed to a moving base of support. Subjects were presented with either low- or high-amplitude broadband visual stimuli during standing posture or while walking on a treadmill at 1 km/h and 5 km/h. Frequency response functions between visual scene motion (input) and trunk kinematics (output) revealed little or no change in the gain of trunk orientation in the standing posture and walking conditions. However, a dramatic increase in gain was observed in trunk (hip and shoulder) horizontal displacement from posture to locomotion. Such increases in gain may be interpreted as an increased coupling to visual scene motion. However, we believe that the increased gain reflects a decrease in stability due to a change of the control problem from standing to locomotion. Indeed, keeping the body upright with the use of vision during walking is complicated by the additional locomotor processes at work. Unlike during standing, vision plays many roles during locomotion, providing information for upright stability as well as body position relative to the external environment.

Surprisingly, despite millions of years of bipedal walking evolution, the gravity-related pendulum mechanism of walking does not seem to be implemented at the onset of independent walking, requiring each toddler to develop it. We discuss the precursor of the mature locomotor pattern in infants as an optimal starting point strategy for gait maturation.

The idea that the CNS may control complex interactions by modular decomposition has received considerable attention. We explored this idea for human locomotion by examining limb kinematics. The coordination of limb segments during human locomotion has been shown to follow a planar law for walking at different speeds, directions, and levels of body unloading. We compared the coordination for different gaits. Eight subjects were asked to walk and run on a treadmill at different speeds or to walk, run, and hop over ground at a preferred speed. To explore various constraints on limb movements, we also recorded stepping over an obstacle, walking with the knees flexed, and air-stepping with body weight support. We found little difference among covariance planes that depended on speed, but there were differences that depended on gait. In each case, we could fit the planar trajectories with a weighted sum of the limb length and orientation trajectories. This suggested that limb length and orientation might provide independent predictors of limb coordination. We tested this further by having the subjects step, run, and hop in place, thereby varying only limb length and maintaining limb orientation fixed, and also by marching with knees locked to maintain limb length constant while varying orientation. The results were consistent with a modular control of limb kinematics where limb movements result from a superposition of separate length- and orientation-related angular covariance. The hypothesis finds support in the animal findings that limb proprioception may also be encoded in terms of these global limb parameters.

Experimental and clinical studies suggest that primate species exhibit greater recovery after lateralized compared to symmetrical spinal cord injuries. Although this observation has major implications for designing clinical trials and translational therapies, advantages in recovery of nonhuman primates over other species has not been shown statistically to date, nor have the associated repair mechanisms been identified. We monitored recovery in more than 400 quadriplegic patients and found that that functional gains increased with the laterality of spinal cord damage. Electrophysiological analyses suggested that corticospinal tract reorganization contributes to the greater recovery after lateralized compared with symmetrical injuries. To investigate underlying mechanisms, we modeled lateralized injuries in rats and monkeys using a lateral hemisection, and compared anatomical and functional outcomes with patients who suffered similar lesions. Standardized assessments revealed that monkeys and humans showed greater recovery of locomotion and hand function than rats. Recovery correlated with the formation of corticospinal detour circuits below the injury, which were extensive in monkeys, but nearly absent in rats. Our results uncover pronounced inter-species differences in the nature and extent of spinal cord repair mechanisms, likely resulting from fundamental differences in the anatomical and functional characteristics of the motor systems in primates versus rodents. Although rodents remain essential for advancing regenerative therapies, the unique response of the primate corticospinal tract after injury re-emphasizes the importance of primate models for designing clinically relevant treatments.

Funding. LG is supported by the EU Project “Integrative approach for the emergence of human like locomotion” (H2R; contract #600698).

International audience; Electrical neuromodulation of lumbar segments improves motor control after spinal cord injury in animal models and humans. However, the physiological principles underlying the effect of this intervention remain poorly understood, which has limited the therapeutic approach to continuous stimulation applied to restricted spinal cord locations. Here we developed stimulation protocols that reproduce the natural dynamics of motoneuron activation during locomotion. For this, we computed the spatiotemporal activation pattern of muscle synergies during locomotion in healthy rats. Computer simulations identified optimal electrode locations to target each synergy through the recruitment of proprioceptive feedback circuits. This framework steered the design of spatially selective spinal implants and real-time control software that modulate extensor and flexor synergies with precise temporal resolution. Spatiotemporal neuromodulation therapies improved gait quality, weight-bearing capacity, endurance and skilled locomotion in several rodent models of spinal cord injury. These new concepts are directly translatable to strategies to improve motor control in humans.

Muscle activity occurring during human locomotion can be accounted for by five basic temporal activation patterns in a variety of locomotion conditions. Here, we examined how these activation patterns interact with muscle activity required for a voluntary movement. Subjects produced a voluntary movement during locomotion, and we examined the resulting kinematics, kinetics, and EMG activity in 16-31 ipsilateral limb and trunk muscles during the tasks. There were four voluntary tasks added to overground walking (approximately 5 km/h) in which subjects kicked a ball, stepped over an obstacle, or reached down and grasped an object on the floor (weight support on either the right or the left foot). Statistical analyses of EMG waveforms showed that the five basic locomotion patterns were invariantly present in each task, although they could be differently weighted across muscles, suggesting a characteristic locomotion timing of muscle activations. We also observed a separate activation that was timed to the voluntary task. The coordination of locomotion with the voluntary task was accomplished by combining activation timings that were associated separately with the voluntary task and locomotion. Activation associated with the voluntary tasks was either synchronous with the timing for locomotion or had additional activations not represented in the basic locomotion timing. We propose that this superposition of an invariant locomotion timing pattern with a voluntary activation timing may be consistent with the proposal suggesting that compound movements are produced through a superposition of motor programs.

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.

Human stepping movements emergein uteroand show several milestones during development to independent walking. Recently, imaging has become an essential tool for investigating the development and function of pattern generation networks in the spinal cord. Here we examine the development of the spinal segmental output by mapping the distribution of motoneuron activity in the lumbosacral spinal cord during stepping in newborns, toddlers, preschoolers, and adults. Newborn stepping is characterized by an alternating bilateral motor output with only two major components that are active at all lumbosacral levels of the spinal cord. This feature was similar across different cycle durations of neonate stepping. The alternating spinal motor output is consistent with a simpler organization of neuronal networks in neonates. Furthermore, a remarkable feature of newborn stepping is a higher overall activation of lumbar versus sacral segments, consistent with a rostrocaudal excitability gradient. In toddlers, the stance-related motor pool activity migrates to the sacral cord segments, while the lumbar motoneurons are separately activated at touchdown. In the adult, the lumbar and sacral patterns become more dissociated with shorter activation times. We conclude that the development of human locomotion from the neonate to the adult starts from a rostrocaudal excitability gradient and involves a gradual functional reorganization of the pattern generation circuitry.

Introduction: Recently, the use of synergy analysis to describe dynamic motor control in cerebral palsy (CP) has become increasingly popular, particularly because synergies have been shown to be a strong predictor of treatment outcome [1]. However, several studies have shown that technical issues such as the matrix factorization algorithms used [2] and EMG processing [3,4] can influence the outcome and interpretation of synergies. Therefore, it is important to study how processing steps can influence synergy outcome measures in CP, especially before and after treatment. Research question: What is the effect of different choices in the EMG analysis (filtering, normalization, and stride selection) on the outcome of synergy analysis in CP, and could this affect the comparison of synergies before and after botulinum neurotoxin (BoNT-A) treatment? Methods: 68 ambulant children with CP (age 9.3 AE 3.6, GMFCS level I–IV) underwent multilevel BoNT-A treatment, and received pre-and 73 AE 25 days post-treatment gait analysis, with EMG measurement of 5 muscles (rectus fem., vastus lat., semitendinosus, tibialis ant. and gastrocnemius med.). EMG data of the most affected leg were high-pass filtered at 20 Hz, rectified, and low-pass filtered at either 2, 3, 10 or 25 Hz. EMG data were either not normalized, normalized to their own maximum, or normalized to the pre-treatment maximum. Finally, EMG was either averaged over all strides (N = 10.8 AE 4.2), or taken from one random stride, or concatenated over all strides. EMG with each combination of filter, normalization, and stride selection was input to a non-negative matrix factorization algorithm (NNMF). An ANOVA for repeated measures was run on the variance accounted for (VAF) by one synergy, to analyse BoNT-A treatment, filtering, normalization, and stride selection effects, as well as their interactions. Results: VAF decreased strongly with increasing filter frequency (Fig. A, p

Regaining Limb Movement Despite many years of intensive research, there is still an urgent need for novel treatments to help patients restore motor function after spinal cord injuries. van den Brand et al. (p. 1182 ) produced left and right hemisections at different levels of the rat thoracic spinal cord to cause complete hind limb paralysis mimicking the situation in humans with spinal cord injury. Systemic application of pharmacological agents, combined with a multisystem rehabilitation program including a robotic postural neuroprosthesis, restored voluntary movements of both hind limbs.

Central nervous system (CNS) disorders distinctly impair locomotor pattern generation and balance, but technical limitations prevent independent assessment and rehabilitation of these subfunctions. Here we introduce a versatile robotic interface to evaluate, enable and train pattern generation and balance independently during natural walking behaviors in rats. In evaluation mode, the robotic interface affords detailed assessments of pattern generation and dynamic equilibrium after spinal cord injury (SCI) and stroke. In enabling mode, the robot acts as a propulsive or postural neuroprosthesis that instantly promotes unexpected locomotor capacities including overground walking after complete SCI, stair climbing following partial SCI and precise paw placement shortly after stroke. In training mode, robot-enabled rehabilitation, epidural electrical stimulation and monoamine agonists reestablish weight-supported locomotion, coordinated steering and balance in rats with a paralyzing SCI. This new robotic technology and associated concepts have broad implications for both assessing and restoring motor functions after CNS disorders, both in animals and in humans. © 2012 Nature America, Inc. All rights reserved.

How rudimentary movements evolve into sophisticated ones during development remains unclear. It is often assumed that the primitive patterns of neural control are suppressed during development, replaced by entirely new patterns. Here we identified the basic patterns of lumbosacral motoneuron activity from multimuscle recordings in stepping neonates, toddlers, preschoolers, and adults. Surprisingly, we found that the two basic patterns of stepping neonates are retained through development, augmented by two new patterns first revealed in toddlers. Markedly similar patterns were observed also in the rat, cat, macaque, and guineafowl, consistent with the hypothesis that, despite substantial phylogenetic distances and morphological differences, locomotion in several animal species is built starting from common primitives, perhaps related to a common ancestral neural network.

The concept of body schema has been introduced and widely discussed in the literature to explain various clinical observations and distortions in the body and space representation. Here we address the role of body schema related information in multi-joint limb motion. The processing of proprioceptive information may differ significantly in static and dynamic conditions since in the latter case the control system may employ specific dynamic rules and constraints. Accordingly, the perception of movement, e.g., estimation of step length and walking distance, may rely on a priori knowledge about intrinsic dynamics of limb segment motion and inherent relationships between gait parameters and body proportions. The findings are discussed in the general framework of space and body movement representation and suggest the existence of a dynamic locomotor body schema used for controlling step length and path estimation.

During the evolution of bipedal modes of locomotion, a sequential rostrocaudal activation of trunk muscles due to the undulatory body movements was replaced by more complex and discrete bursts of activity. Nevertheless, the capacity for segmental rhythmogenesis and the rostrocaudal propagation of spinal cord activity has been conserved. In humans, motoneurons of different muscles are arranged in columns, with a specific grouping of muscles at any given segmental level. The muscle patterns of locomotor activity and the biomechanics of the body center of mass have been studied extensively, but their interrelationship remains poorly understood. Here we mapped the electromyographic activity recorded from 30 bilateral leg muscles onto the spinal cord in approximate rostrocaudal locations of the motoneuron pools during walking and running in humans. We found that the rostrocaudal displacements of the center of bilateral motoneuron activity mirrored the changes in the energy due to the center-of-body mass motion. The results suggest that biomechanical mechanisms of locomotion, such as the inverted pendulum in walking and the pogo-stick bouncing in running, may be tightly correlated with specific modes of progression of motor pool activity rostrocaudally in the spinal cord.

In adults, locomotor movements are accommodated to various support surface conditions by means of specific anticipatory locomotor adjustments and changes in the intersegmental coordination. Here we studied the kinematic strategies of toddlers at the onset of independent walking when negotiating various support surface conditions: stepping over an obstacle, walking on an inclined surface, and on a staircase. Generally, toddlers could perform these tasks only when supported by the arm. They exhibited strategies very different from those of the adults. Although adults maintained walking speed roughly constant, toddlers markedly accelerated when walking downhill or downstairs and decelerated when walking uphill or upstairs. Their coordination pattern of thigh–shank–foot elevation angles exhibited greater inter-trial variability than that in adults, but it did not undergo the systematic change as a function of task that was present in adults. Thus the intersegmental covariance plane rotated across tasks in adults, whereas its orientation remained roughly constant in toddlers. In contrast with the adults, the toddlers often tended to place the foot onto the obstacle or across the edges of the stairs. We interpret such foot placements as part of a haptic exploratory repertoire and we argue that the maintenance of a roughly constant planar covariance—irrespective of the surface inclination and height—may be functional to the exploratory behavior. The latter notion is consistent with the hypothesis proposed decades ago by Bernstein that, when humans start to learn a skill, they may restrict the number of degrees of freedom to reduce the size of the search space and simplify the coordination.

Objective. Decoding forelimb movements from the firing activity of cortical neurons has been interfaced with robotic and prosthetic systems to replace lost upper limb functions in humans. Despite the potential of this approach to improve locomotion and facilitate gait rehabilitation, decoding lower limb movement from the motor cortex has received comparatively little attention. Here, we performed experiments to identify the type and amount of information that can be decoded from neuronal ensemble activity in the hindlimb area of the rat motor cortex during bipedal locomotor tasks. Approach. Rats were trained to stand, step on a treadmill, walk overground and climb staircases in a bipedal posture. To impose this gait, the rats were secured in a robotic interface that provided support against the direction of gravity and in the mediolateral direction, but behaved transparently in the forward direction. After completion of training, rats were chronically implanted with a micro-wire array spanning the left hindlimb motor cortex to record single and multi-unit activity, and bipolar electrodes into 10 muscles of the right hindlimb to monitor electromyographic signals. Whole-body kinematics, muscle activity, and neural signals were simultaneously recorded during execution of the trained tasks over multiple days of testing. Hindlimb kinematics, muscle activity, gait phases, and locomotor tasks were decoded using offline classification algorithms. Main results. We found that the stance and swing phases of gait and the locomotor tasks were detected with accuracies as robust as 90% in all rats. Decoded hindlimb kinematics and muscle activity exhibited a larger variability across rats and tasks. Significance. Our study shows that the rodent motor cortex contains useful information for lower limb neuroprosthetic development. However, brain-machine interfaces estimating gait phases or locomotor behaviors, instead of continuous variables such as limb joint positions or speeds, are likely to provide more robust control strategies for the design of such neuroprostheses.

The inverted pendulum model in which the centre of mass of the body vaults over the stance leg in an arc represents a basic mechanism of bipedal walking. Is the pendulum mechanism innate, or is it learnt through walking experience? We studied eight toddlers (about 1 year old) at their first unsupported steps, 18 older children (1.3-13 years old), and ten adults. Two infants were also tested repeatedly over a period of 4 months before the onset of independent walking. Pendulum mechanism was quantified from the kinematics of the greater trochanter, correlation between kinetic and gravitational potential energy of the centre of body mass obtained from the force plate recordings, and percentage of recovery of mechanical energy. In toddlers, these parameters deviated significantly (P, Comparative Study, Journal Article, Research Support, Non-U.S. Gov't, info:eu-repo/semantics/published