Sequential activation of axial muscles during different forms of rhythmic behavior in man

In humans, studies of back muscle activity have mainly addressed the functioning of lumbar muscles during postural adjustments or rhythmic activity, including locomotor tasks. The present study investigated how back muscles are activated along the spine during rhythmical activities in order to gain insights into spinal neuronal organization. Electromyographic recordings of back muscles were performed at various trunk levels, and changes occurring in burst amplitudes and phase relationships were analyzed. Subjects performed several rhythmic behaviors: forward walking (FW), backward walking (BW), amble walking (where the subjects moved their arms in phase with the ipsilateral leg), walking on hands and knees (HK) and walking on hands with the knees on the edge of a treadmill (Hand). In a final task, the subjects were standing and were asked to swing (Swing) only their arms as if they were walking. It was found that axial trunk muscles are sequentially activated by a motor command running along the spinal cord (which we term “motor waves”) during various types of locomotion or other rhythmic motor tasks. The bursting pattern recorded under these conditions can be classified into three categories: (1) double-burst rhythmic activity in a descending (i.e., with a rostro-caudal propagation) motor wave during FW, BW and HK conditions; (2) double-burst rhythmic activity with a stationary motor wave (i.e., occurring in a single phase along the trunk) during the ‘amble’ walk condition; (3) monophasic rhythmic activity with an ascending (i.e., with a caudo-rostral propagation) motor wave during the Swing and Hands conditions. Our results suggest that the networks responsible for the axial propagation of motor activity during locomotion may correspond to those observed in invertebrates or lower vertebrates, and thus may have been partly phylogenetically conserved. Such an organization could support the dynamic control of posture by ensuring fluent movement during locomotion.     

Predictive Simulations of Neuromuscular Coordination and Joint-Contact Loading in Human Gait

We implemented direct collocation on a full-body neuromusculoskeletal model to calculate muscle forces, ground reaction forces and knee contact loading simultaneously for one cycle of human gait. A data-tracking collocation problem was solved for walking at the normal speed to establish the practicality of incorporating a 3D model of articular contact and a model of foot–ground interaction explicitly in a dynamic optimization simulation. The data-tracking solution then was used as an initial guess to solve predictive collocation problems, where novel patterns of movement were generated for walking at slow and fast speeds, independent of experimental data. The data-tracking solutions accurately reproduced joint motion, ground forces and knee contact loads measured for two total knee arthroplasty patients walking at their preferred speeds. RMS errors in joint kinematics were?<?2.0° for rotations and?<?0.3 cm for translations while errors in the model-computed ground-reaction and knee-contact forces were?<?0.07 BW and?<?0.4 BW, respectively. The predictive solutions were also consistent with joint kinematics, ground forces, knee contact loads and muscle activation patterns measured for slow and fast walking. The results demonstrate the feasibility of performing computationally-efficient, predictive, dynamic optimization simulations of movement using full-body, muscle-actuated models with realistic representations of joint function.     

Forward and backward arm cycling are regulated by equivalent neural mechanisms

It was shown some time ago that cutaneous reflexes were phase-reversed when comparing forward and backward treadmill walking. Activity of central-pattern-generating networks (CPG) regulating neural activity for locomotion was suggested as a mechanism involved in this "program reversal." We have been investigating the neural control of arm movements and the role for CPG mechanisms in regulating rhythmic arm cycling. The purpose of this study was to evaluate the pattern of muscle activity and reflex modulation when comparing forward and backward arm cycling. During rhythmic arm cycling (forward and backward), cutaneous reflexes were evoked with trains (5 x 1.0 ms pulses at 300 Hz) of electrical stimulation delivered to the superficial radial (SR) nerve at the wrist. Electromyographic (EMG) recordings were made bilaterally from muscles acting at the shoulder, elbow, and wrist. Analysis was conducted on specific sections of the movement cycle after phase-averaging contingent on the timing of stimulation in the movement cycle. EMG patterns for rhythmic arm cycling are similar during both forward and backward motion. Cutaneous reflex amplitudes were similarly modulated at both early and middle latency irrespective of arm cycling direction. That is, at similar phases in the movement cycle, responses of corresponding sign and amplitude were seen regardless of movement direction. The results are generally parallel to the observations seen in leg muscles after stimulation of cutaneous nerves in the foot during forward and backward walking and provide further evidence for CPG activity contributing to neural activation and reflex modulation during rhythmic arm movement.     

How is the normal locomotor program modified to produce backward walking?

Summary The modifications occurring in the movement and muscle activity patterns of the leg when changing from forward to backward walking were studied in five healthy subjects during walking on a motor driven treadmill. Movements were recorded with a Selspot optoelectronic system and muscle activity with electromyography using surface electrodes. The movement trajectories of the leg in forward and backward walking essentially mirrored each other, even though the movements occurred in the reversed direction. The angular displacements at the hip, knee and ankle joints showed similar overall magnitude and pattern in the two situations. Most of the investigated muscles changed their pattern of activity in relation to the different movement phases. At the ankle, there was a switch between flexors and extensors with flexor activation during support in backward walking. The bursts of activity in knee extensors were prolonged and shifted to the main part of the support phase. In the hip extensors, the activity periods retained their positions relative to the leg movements, but changed function due to the reversed direction of movement. Thus, drastic changes occur in the normal locomotor program to produce a reversal of leg movements and propulsion backwards.     

Motor patterns in human walking and running

Despite distinct differences between walking and running, the two types of human locomotion are likely to be controlled by shared pattern-generating networks. However, the differences between their kinematics and kinetics imply that corresponding muscle activations may also be quite different. We examined the differences between walking and running by recording kinematics and electromyographic (EMG) activity in 32 ipsilateral limb and trunk muscles during human locomotion, and compared the effects of speed (3-12 km/h) and gait. We found that the timing of muscle activation was accounted for by five basic temporal activation components during running as we previously found for walking. Each component was loaded on similar sets of leg muscles in both gaits but generally on different sets of upper trunk and shoulder muscles. The major difference between walking and running was that one temporal component, occurring during stance, was shifted to an earlier phase in the step cycle during running. These muscle activation differences between gaits did not simply depend on locomotion speed as shown by recordings during each gait over the same range of speeds (5-9 km/h). The results are consistent with an organization of locomotion motor programs having two parts, one that organizes muscle activation during swing and another during stance and the transition to swing. The timing shift between walking and running reflects therefore the difference in the relative duration of the stance phase in the two gaits.     

Detecting temporal reversals in human locomotion

An experiment investigated the ability by human observers to detect temporal reversals in dynamic displays of human locomotion. We video-taped the lower portion of the body of actors walking at their preferred speed either in the normal, forward direction (FW) or in the backward direction (BW). The videos were presented in a random order either as recorded (N) or in reverse (R). In one session, we presented both normal and time-reversed stimuli in the original upright orientation. In a second session, the stimuli were rotated by 180° around the horizontal axis. Observers were informed that the real recorded movement was either forward or backward and were asked to decide whether or not the movement had been time-reversed prior to the presentation. Although the kinematics of forward and backward human locomotion is quite similar, the detection of temporal reversals followed a consistent pattern showing a good accuracy in condition FW-N and a reduced but still above-chance performance in condition BW-R (by design, in both conditions actors appeared to walk forward). Performance was instead at chance level in the other two conditions where the apparent direction of the movement was backward. Inverting the spatial orientation of the stimuli reduced but did not suppress the ability to detect temporal reversals in the two conditions with apparent forward direction of movement. It is argued that implicit motor competence is at least in part instrumental for extracting the subtle discriminal information from the stimuli.     

The effects of human ankle muscle vibration on posture and balance during adaptive locomotion

This study investigated the contribution of ankle muscle proprioception to the control of dynamic stability and lower limb kinematics during adaptive locomotion, by using mechanical vibration to alter the muscle spindle output of individuals' stance limbs. It was hypothesised that muscle length information from the ankle of the stance limb provides information describing location as well as acceleration of the centre of mass (COM) with respect to the support foot during the swing phase of locomotion. Our prediction, based on this hypothesis was that ankle muscle vibration would cause changes to the position and acceleration of the COM and/or compensatory postural responses. Vibrators were attached to both the stance limb ankle plantarflexors (at the Achilles tendon) and the opposing dorsiflexor muscle group (over tibialis anterior). Participants were required to walk along a 9-m travel path and step over any obstacles placed in their way. There were three task conditions: (1) an obstacle (15 cm in height) was positioned at the midpoint of the walkway prior to the start of the trial, (2) the same obstacle was triggered to appear unexpectedly one step in front of the participant at the walkway midpoint and (3) the subjects' walking path remained clear. The participants' starting position was manipulated so that the first step over the obstacle (when present) was always performed with their right leg. For each obstacle condition participants experienced the following vibration conditions: no vibration, vibration of the left leg calf muscles or vibration of the anterior compartment muscles of the lower left leg. Vibration began one step before the obstacle at left leg heel contact and continued for 1 s. Vibrating the ankle muscles of the stance limb during the step over an obstacle resulted in significant changes to COM behaviour [measured as displacement, acceleration and position with respect to the centre of pressure (COP)] in both the medial/lateral (M/L) and anterior/posterior planes. There were also significant task-specific changes in stepping behaviour associated with COM control (measured as peak M/L acceleration, M/L foot displacement and COP position under the stance foot during the step over the obstacle). The results provide strong evidence that the primary endings of ankle muscle spindles play a significant role in the control of posture and balance during the swing phase of locomotion by providing information describing the movement of the body's COM with respect to the support foot. Our results also provide supporting evidence for the proposal that there are context-dependent changes in muscle spindle sensitivity during human locomotion.     

Activity of motor cortex neurons during backward locomotion

Forward walking (FW) and backward walking (BW) are two important forms of locomotion in quadrupeds. Participation of the motor cortex in the control of FW has been intensively studied, whereas cortical activity during BW has never been investigated. The aim of this study was to analyze locomotion-related activity of the motor cortex during BW and compare it with that during FW. For this purpose, we recorded activity of individual neurons in the cat during BW and FW. We found that the discharge frequency in almost all neurons was modulated in the rhythm of stepping during both FW and BW. However, the modulation patterns during BW and FW were different in 80% of neurons. To determine the source of modulating influences (forelimb controllers vs. hindlimb controllers), the neurons were recorded not only during quadrupedal locomotion but also during bipedal locomotion (with either forelimbs or hindlimbs walking), and their modulation patterns were compared. We found that during BW (like during FW), modulation in some neurons was determined by inputs from limb controllers of only one girdle, whereas the other neurons received inputs from both girdles. The combinations of inputs could depend on the direction of locomotion. Most often (in 51% of forelimb-related neurons and in 34% of the hindlimb-related neurons), the neurons received inputs only from their own girdle when this girdle was leading and from both girdles when this girdle was trailing. This reconfiguration of inputs suggests flexibility of the functional roles of individual cortical neurons during different forms of locomotion.     

Balance control in patients with distal versus proximal muscle weakness

Muscle weakness is consistently associated with falls in the elderly people, typically when present along with other risk factors. However, it remains unknown whether and how muscle weakness alone affects balance. This hampers development of more effective fall prevention strategies. Clinical observations suggest that the amount and distribution of muscle weakness influences balance control. We therefore investigated balance corrections in patients with either predominantly proximal (limb girdle muscular dystrophy (LGMD); n=8) or distal (distal spinal muscular atrophy; n=5) leg weakness, and 27 matched healthy controls. Balance was perturbed using surface tilt rotations that were delivered randomly in eight directions. Balance measures were full body kinematics and surface electromyographic activity (EMG) of leg, arm, and trunk muscles. Both patient groups were more unstable than controls, as reflected by greater excursions of the centre of mass (COM), especially in the pitch (anterior–posterior (AP)) plane. COM displacements were greater in distal weakness patients. Patients with distal weakness had excessive and unstable trunk, knee and ankle movements, and this was present following both forward and backward directed balance perturbations, possibly reflecting the greater use of distal leg muscles in these directions. In contrast, the less weak proximal weakness patients demonstrated unstable trunk and ankle movements only for backward directed balance perturbations. Both patient groups used arm movements to compensate for their instability. We conclude that primarily distal but also proximal muscle weakness leads to significant postural instability. This observation, together with the retained ability of patients to use compensatory arm movements, provides targets that may be amenable to improvement with therapeutic intervention.     

Phase-dependent reflex reversal in human leg muscles during walking

1. Reflex responses during walking were elicited in humans by stimulation of the tibial nerve at the ankle. The stimulus intensity was controlled by monitoring the M-wave from an intrinsic foot muscle. Responses were observed in the ipsilateral tibialis anterior (TA), soleus (SO), and rectus femoris (RF) muscles. The most reproducible responses were observed at a middle latency between 50 and 90 ms. The responses were most likely of cutaneous origin, because they closely resembled the responses to stimulation of a purely cutaneous nerve, the sural nerve. 2. A reversal in the direction of the middle latency response from excitation to inhibition was observed for the first time within single muscles during walking. Evidence for a reversal was seen in all three muscles examined and in all seven subjects. 3. The reflex reversal could not be elicited in standing. An inhibition whose amplitude varied in a linear fashion with stimulus intensity and background activation level was always observed at middle latency. The responses elicited during standing resembled those during the stance phase of walking. The two tasks shared some common movement goals and appeared to make use of similar reflex pathways.     

Control of reflex reversal in stick insect walking: effects of intersegmental signals, changes in direction, and optomotor-induced turning

In many animals, the effects of sensory feedback on motor output change during locomotion. These changes can occur as reflex reversals in which sense organs that activate muscles to counter perturbations in posture control instead reinforce movements in walking. The mechanisms underlying these changes are only partially understood. As such, it is unclear whether reflex reversals are modulated when locomotion is adapted, such as during changes in walking direction or in turning movements. We investigated these questions in the stick insect Carausius morosus, where sensory signals from the femoral chordotonal organ are known to produce resistance reflexes at rest but assistive movements during walking. We studied how intersegmental signals from neighboring legs affect the generation of reflex reversals in a semi-intact preparation that allows free leg movement during walking. We found that reflex reversal was enhanced by stepping activity of the ipsilateral neighboring rostral leg, whereas stepping of contralateral legs had no effect. Furthermore, we found that the occurrence of reflex reversals was task-specific: in the front legs of animals with five legs walking, reflex reversal was generated only during forward and not backward walking. Similarly, during optomotor-induced curved walking, reflex reversal occurred only in the middle leg on the inside of the turn and not in the contralateral leg on the outside of the turn. Thus our results show for the first time that the nervous system modulates reflexes in individual legs in the adaptation of walking to specific tasks.     

Neural compensation within the human triceps surae during prolonged walking

During human walking, muscle activation strategies are approximately constant across consecutive steps over a short time, but it is unknown whether they are maintained over a longer duration. Prolonged walking may increase tendinous tissue (TT) compliance, which can influence neural activation, but the neural responses of individual muscles have not been investigated. This study investigated the hypothesis that muscle activity is up- or down-regulated in individual triceps surae muscles during prolonged walking. Thirteen healthy subjects walked on a treadmill for 60 min at 4.5 km/h, while triceps surae muscle activity, maximal muscle compound action potentials, and kinematics were recorded every 5 min, and fascicle lengths were estimated at the beginning and end of the protocol using ultrasound. After 1 h of walking, soleus activity increased by 9.3 ± 0.2% (P < 0.05) and medial gastrocnemius activity decreased by 9.3 ± 0.3% (P < 0.01). Gastrocnemius fascicle length at ground contact shortened by 4.45 ± 0.99% (P < 0.001), whereas soleus fascicle length was unchanged (P = 0.988). Throughout the stance phase, medial gastrocnemius fascicle lengthening decreased by 44 ± 13% (P < 0.001), whereas soleus fascicle lengthening amplitude was unchanged (P = 0.650). The data suggest that a compensatory neural strategy exists between triceps surae muscles and that changes in muscle activation are generally mirrored by changes in muscle fascicle length. These findings also support the notion of muscle-specific changes in TT compliance after prolonged walking and highlight the ability of the CNS to maintain relatively constant movement patterns in spite of neuromechanical changes in individual muscles.     

Coordinated modulation of locomotor muscle synergies constructs straight-ahead and curvilinear walking in humans

We describe the muscle synergies accompanying steering of walking along curved trajectories, in order to analyze the simultaneous control of progression and balance-threatening emerging forces. For this purpose, we bilaterally recorded in ten subjects the electromyograms (EMGs) of a representative sample of leg and trunk muscles (n=16) during continuous walking along one straight and two curved trajectories at natural speed. Curvilinear locomotion involved a graded, limb-dependent modulation of amplitude and timing of activity of the muscles of the legs and trunk. The turn-related modulation of the motor pattern was highly coordinated amongst muscles and body sides. For all muscles, linear relationships were detected between the spatial and temporal features of muscle EMG activity. The largest modulation of EMG was observed in gastrocnemius medialis and lateralis muscles, which showed opposite changes in timing and amplitude during curve-walking. Moreover, amplitude and timing characteristics of muscle activities were significantly correlated with the spatial and temporal gait adaptations that are associated with curvilinear locomotion. The present results reveal that fine-modulation of the muscle synergies underlying straight-ahead locomotion is enough to generate the adequate propulsive forces to steer walking and maintain balance. These findings suggest that the turn-related command operates by modulation of the phase relationships between the tightly coupled neuronal assemblies that drive motor neuron activity during walking. This would produce the invariant templates for locomotion kinematics that are at the base of human navigation in space.     

Coordination of intrinsic and extrinsic foot muscles during walking

Purpose

The human foot undergoes complex deformations during walking due to passive tissues and active muscles. However, based on prior recordings it is unclear if muscles that contribute to flexion/extension of the metatarsophalangeal (MTP) joints are activated synchronously to modulate joint impedance, or sequentially to perform distinct biomechanical functions. We investigated the coordination of MTP flexors and extensors with respect to each other, and to other ankle–foot muscles.

Methods

We analyzed surface electromyographic (EMG) recordings of intrinsic and extrinsic foot muscles for healthy individuals during level treadmill walking, and also during sideways and tiptoe gaits. We computed stride-averaged EMG envelopes and used the timing of peak muscle activity to assess synchronous vs. sequential coordination.

Results

We found that peak MTP flexor activity occurred significantly before peak MTP extensor activity during walking (P < 0.001). The period around stance-to-swing transition could be roughly characterized by sequential peak muscle activity from the ankle plantarflexors, MTP flexors, MTP extensors, and then ankle dorsiflexors. We found that foot muscles that activated synchronously during forward walking tended to dissociate during other locomotor tasks. For instance, extensor hallucis brevis and extensor digitorum brevis muscle activation peaks decoupled during sideways gait.

Conclusions

The sequential peak activity of MTP flexors followed by MTP extensors suggests that their biomechanical contributions may be largely separable from each other and from other extrinsic foot muscles during walking. Meanwhile, the task-specific coordination of the foot muscles during other modes of locomotion indicates a high-level of specificity in their function and control.

     

Informational and Neuromuscular Contributions to Anchoring in Rhythmic Wrist Cycling

Continuous rhythmic movements are often geared toward particular points in the movement cycle, as evidenced by a local reduction in trajectory variability. These so-called anchor points provide a window into motor control, since changes in the degree of anchoring may reveal how informational and/or neuromuscular properties are exploited in the organization of rhythmic movements. The present experiment examined the relative contributions of informational timing (metronome beeps) and neuromuscular (wrist postures) constraints on anchoring by systematically varying both factors at movement reversal points. To this end, participants cycled their right wrist in a flexed, neutral, or extended posture, either self-paced or synchronized to a metronome pacing peak flexion, peak extension, or both peak flexion and extension. The effects of these manipulations were assessed in terms of kinematics, auditory-motor coordination, and muscle activity. The degree of anchoring seen at the reversal points depended on the degree of compatibility of the prevailing configuration of neuromuscular and informational timing constraints, which had largely independent effects. We further observed systematic changes in muscular activity, which revealed distinct contributions of posture- and muscle-dependent neuromuscular properties to motor control. These findings indicate that the anchor-based discretization of the control of continuous rhythmic wrist movements is determined by both informational timing and neuromuscular constraints in a task-specific manner with subtle interactions between the two, and exemplify how movement variability may be exploited to gain such insights.     

Musculoskeletal modelling of the Nile crocodile (Crocodylus niloticus) hindlimb: Effects of limb posture on leverage during terrestrial locomotion

We developed a three-dimensional, computational biomechanical model of a juvenile Nile crocodile (Crocodylus niloticus) pelvis and hindlimb, composed of 47 pelvic limb muscles, to investigate muscle function. We tested whether crocodiles, which are known to use a variety of limb postures during movement, use limb orientations (joint angles) that optimise the moment arms (leverages) or moment-generating capacities of their muscles during different limb postures ranging from a high walk to a sprawling motion. We also describe the three-dimensional (3D) kinematics of the crocodylian hindlimb during terrestrial locomotion across an instrumented walkway and a treadmill captured via X-ray Reconstruction of Moving Morphology (biplanar fluoroscopy; ‘XROMM’). We reconstructed the 3D positions and orientations of each of the hindlimb bones and used dissection data for muscle lines of action to reconstruct a focal, subject-specific 3D musculoskeletal model. Motion data for different styles of walking (a high, crouched, bended and two types of sprawling motion) were fed into the 3D model to identify whether any joints adopted near-optimal poses for leverage across each of the behaviours. We found that (1) the hip adductors and knee extensors had their largest leverages during sprawling postures and (2) more erect postures typically involved greater peak moment arms about the hip (flexion-extension), knee (flexion) and metatarsophalangeal (flexion) joints. The results did not fully support the hypothesis that optimal poses are present during different locomotory behaviours because the peak capacities were not always reached around mid-stance phase. Furthermore, we obtained few clear trends for isometric moment-generating capacities. Therefore, perhaps peak muscular leverage in Nile crocodiles is instead reached either in early/late stance or possibly during swing phase or other locomotory behaviours that were not studied here, such as non-terrestrial movement. Alternatively, our findings could reflect a trade-off between having to execute different postures, meaning that hindlimb muscle leverage is not optimised for any singular posture or behaviour. Our model, however, provides a comprehensive set of 3D estimates of muscle actions in extant crocodiles which can form a basis for investigating muscle function in extinct archosaurs.     

A comparison of the effects of agonist and antagonist muscle fatigue on performance of rapid movements

The aim of this study was to investigate the effects of agonist and antagonist muscle fatigue on the performance of rapid, self-terminating movements. Six subjects performed rapid, consecutive elbow flexion and extension movements between two targets prior to and after fatiguing either the elbow flexor or elbow extensor muscles. The experiments demonstrated consistent results. Agonist muscle fatigue was associated with a decrease in peak velocity and peak deceleration, while a decrease in peak acceleration was particularly prominent. Antagonist muscle fatigue, however, was associated with a decrease in peak deceleration, while a decrease in both the peak velocity and peak acceleration was modest and, in some tests, non-significant. The relative acceleration time (i.e. acceleration time as a proportion of the total movement time) increased when agonists were fatigued, but decreased when antagonists were fatigued. Taken together, these results emphasize the mechanical roles of the agonist and antagonist muscles; namely, the fatigue of each muscle group particularly affected the movement phase in which that group accelerated a limb, while changes of the movement kinematics pattern provided more time for action of the fatigued muscles. In addition, the results presented suggest that agonist muscle fatigue affects movement velocity more than antagonist muscle fatigue, even in movements that demonstrate prominently both mechanical and myoelectric activity of the antagonist muscles, such as rapid, self-terminating movements.     

Recumbent stepping has similar but simpler neural control compared to walking

The purpose of this study was to compare muscle activation patterns and kinematics during recumbent stepping and walking to determine if recumbent stepping has a similar motor pattern as walking. We measured joint kinematics and electromyography in ten neurologically intact humans walking on a treadmill at 0 and 50% body weight support (BWS), and recumbent stepping using a commercially available exercise machine. Cross correlation of upper and lower limb electromyography patterns between conditions revealed high correlations for most muscles. A principal component analysis revealed that the first factor accounted for more muscle activation signal content during recumbent stepping (81%) than during walking (70%). This indicates that the motor pattern during walking is more complex than during stepping. Cross correlation analysis found a high correlation between factors for recumbent stepping and walking (R = 0.54), though not as high as the correlation between factors for walking at 0% BWS and walking at 50% BWS (R = 0.68). There were substantial differences in joint kinematics between walking and recumbent stepping, most notably in hip, elbow, and shoulder motions. These results suggest that although the two tasks have different kinematic patterns, recumbent stepping relies on similar neural networks as walking. Individuals with neurological impairments may be able to improve walking ability from recumbent stepping practice given similarities in neural control between the two tasks.     

Automatic vs hand-controlled walking of paraplegics

A rule-based control and its application in functional electrical stimulation (FES) assisted walking of subjects with paraplegia are described in this paper. The design of rules for control comprises the following two steps: (1) determination of muscle activation patterns by using a fully customized spatial (3D) model of paraplegic walking, and (2) learning of rules, that is, correlation between the muscle activation patterns and kinematics of walking by means of an artificial neural network. The adopted FES system activated eight muscle groups with surface electrodes. The only joints allowing movement in the coronal plane were the hips, and externally controlled joints in sagittal plane were ankles, knees and hips. The simulation minimized the tracking error of the joint angles and the total activation of all eight muscles being stimulated. A radial-basis function artificial neural network was applied for learning of rules. Three automatically controlled modes (slow, near-normal, and near-ballistic) and hand-controlled walking were evaluated in six subjects with a complete spinal cord lesion (T8-T10). The performance of walking was assessed by the following: (1) energy consumption based on oxygen uptake, (2) physiological cost index, (3) maximum speed of walking, and (4) a questionnaire. The results showed that all modes of walking are achievable and that automatic control leads to more efficient and faster walking. The speed of walking achieved by automatic control was almost three times bigger compared with the speed of hand-controlled walking. The energy cost and rate decreased significantly when automatic control was applied; yet, they were still much bigger than the values measured in able-bodied subjects. The objective outcome measures suggest that the near-ballistic walking was the most effective, yet a questionnaire shows that most subjects preferred slow walking. The most likely reason for the preference of lower efficiency walking over the faster end energy efficient near-ballistic walking was that paraplegic patients had difficulties in synchronizing the voluntary movement of the trunk and arms to the artificially controlled movements of legs.     

The up and down bobbing of human walking: a compromise between muscle work and efficiency

Human walking has a peculiar straight-legged style. Consequently, the body's centre of mass (CM) moves up and down with each step, which is noticeable in their up and down head bobbing while walking. This vertical CM movement enables humans to save energy via a pendulum-like mechanism but is probably a relatively recent locomotor innovation insofar as earliest bipeds may have walked flexed and flat. We investigated the mechanics, energetics, muscle efficiency and optimization of human walking by decreasing and increasing the vertical CM displacement (flat and bouncy walking) in comparison to normal walking at six speeds (1–6 km h⁻¹). In both flat and bouncy walking, the pendular mechanism was reduced and the energy cost was increased. However, this increase was unexpectedly much sharper in flat walking where muscles provided normal mechanical work but with a decrease in muscle efficiency. In bouncy walking, muscles provided extra mechanical work in an efficient way. Our results showed that not only do humans bob up and down in normal walking to save energy via a pendulum-like mechanism but also to make their muscles work efficiently. Actually, walking flat makes the muscles work in unfavourable conditions that waste energy. Furthermore, we are still close to a flat CM displacement relative to our current ability to change this displacement, which suggests that reducing vertical CM displacement is indeed important but only to certain limits. Evolution may ultimately have chosen the best compromise between flat locomotion that requires little work to move and bouncy locomotion that improves muscle efficiency to minimize energy consumption.