Light Accelerates Morphogenesis and Acquisition of Interlimb Stepping in Chick Embryos

Chicks are bipedal precocious vertebrates that achieve adaptive locomotor skill within hours after hatching. Development of limb movement has been extensively studied in the chicken embryo, but few studies have focused on the preparations leading to precocious locomotor skill. Chicks typically hatch after 21 days of incubation, and recent studies provided evidence that the neural circuits for intralimb control of stepping are established between embryonic days (E) 18–20. It has also been shown that variations in light exposure during embryogenesis can accelerate or delay the onset of hatching and walking by 1 to 2 days. Our earlier work revealed that despite these differences in time to hatch, chicks incubated in different light conditions achieved similar locomotor skill on the day of hatching. Results suggested to us that light exposure during incubation may have accelerated development of locomotor circuits in register with earlier hatching. Thus, in this study, embryos were incubated in 1 of 3 light conditions to determine if development of interlimb coordination at a common time point, 19 days of incubation, varied with light exposure during embryogenesis. Leg muscle activity was recorded bilaterally and burst analyses were performed for sequences of spontaneous locomotor-related activity in one or more ankle muscles to quantify the extent of interlimb coordination in ovo. We report findings indicating that the extent of interlimb coordination varied with light exposure, and left-right alternating steps were a more reliable attribute of interlimb coordination for embryos incubated in constant bright light. We provide evidence that morphological development of the leg varied with light exposure. Based on these findings, we propose that light can accelerate the development of interlimb coordination in register with earlier hatching. Our results lead us to further propose that alternating left-right stepping is the default pattern of interlimb coordination produced by locomotor circuits during embryogenesis.     

The metabolic cost of changing walking speeds is significant,implies lower optimal speeds for shorter distances,and increases daily energy estimates

Humans do not generally walk at constant speed, except perhaps on a treadmill. Normal walking involves starting, stopping and changing speeds, in addition to roughly steady locomotion. Here, we measure the metabolic energy cost of walking when changing speed. Subjects (healthy adults) walked with oscillating speeds on a constant-speed treadmill, alternating between walking slower and faster than the treadmill belt, moving back and forth in the laboratory frame. The metabolic rate for oscillating-speed walking was significantly higher than that for constant-speed walking (6–20% cost increase for ±0.13–0.27 m s⁻¹ speed fluctuations). The metabolic rate increase was correlated with two models: a model based on kinetic energy fluctuations and an inverted pendulum walking model, optimized for oscillating-speed constraints. The cost of changing speeds may have behavioural implications: we predicted that the energy-optimal walking speed is lower for shorter distances. We measured preferred human walking speeds for different walking distances and found people preferred lower walking speeds for shorter distances as predicted. Further, analysing published daily walking-bout distributions, we estimate that the cost of changing speeds is 4–8% of daily walking energy budget.     

Walking speed changes in response to novel user-driven treadmill control

Implementing user-driven treadmill control in gait training programs for rehabilitation may be an effective means of enhancing motor learning and improving functional performance. This study aimed to determine the effect of a user-driven treadmill control scheme on walking speeds, anterior ground reaction forces (AGRF), and trailing limb angles (TLA) of healthy adults. Twenty-three participants completed a 10-m overground walking task to measure their overground self-selected (SS) walking speeds. Then, they walked at their SS and fastest comfortable walking speeds on an instrumented split-belt treadmill in its fixed speed and user-driven control modes. The user-driven treadmill controller combined inertial-force, gait parameter, and position based control to adjust the treadmill belt speed in real time. Walking speeds, peak AGRF, and TLA were compared among test conditions using paired t-tests (α = 0.05). Participants chose significantly faster SS and fast walking speeds in the user-driven mode than the fixed speed mode (p > 0.05). There was no significant difference between the overground SS walking speed and the SS speed from the user-driven trials (p < 0.05). Changes in AGRF and TLA were caused primarily by changes in walking speed, not the treadmill controller. Our findings show the user-driven treadmill controller allowed participants to select walking speeds faster than their chosen speeds on the fixed speed treadmill and similar to their overground speeds. Since user-driven treadmill walking increases cognitive activity and natural mobility, these results suggest user-driven treadmill control would be a beneficial addition to current gait training programs for rehabilitation.     

Running for Exercise Mitigates Age-Related Deterioration of Walking Economy

Introduction

Impaired walking performance is a key predictor of morbidity among older adults. A distinctive characteristic of impaired walking performance among older adults is a greater metabolic cost (worse economy) compared to young adults. However, older adults who consistently run have been shown to retain a similar running economy as young runners. Unfortunately, those running studies did not measure the metabolic cost of walking. Thus, it is unclear if running exercise can prevent the deterioration of walking economy.

Purpose

To determine if and how regular walking vs. running exercise affects the economy of locomotion in older adults.

Methods

15 older adults (69±3 years) who walk ≥30 min, 3x/week for exercise, “walkers” and 15 older adults (69±5 years) who run ≥30 min, 3x/week, “runners” walked on a force-instrumented treadmill at three speeds (0.75, 1.25, and 1.75 m/s). We determined walking economy using expired gas analysis and walking mechanics via ground reaction forces during the last 2 minutes of each 5 minute trial. We compared walking economy between the two groups and to non-aerobically trained young and older adults from a prior study.

Results

Older runners had a 7–10% better walking economy than older walkers over the range of speeds tested (p = .016) and had walking economy similar to young sedentary adults over a similar range of speeds (p = .237). We found no substantial biomechanical differences between older walkers and runners. In contrast to older runners, older walkers had similar walking economy as older sedentary adults (p = .461) and ∼26% worse walking economy than young adults (p<.0001).

Conclusion

Running mitigates the age-related deterioration of walking economy whereas walking for exercise appears to have minimal effect on the age-related deterioration in walking economy.     

Infant stepping: a window to the behaviour of the human pattern generator for walking

The aim of this paper is to provide evidence, both published and new, to support the notion that human infants are particularly good subjects for the study of the pattern generator for walking. We and others have shown that stepping can be initiated by sensory input from the legs or by general heightened excitability of the infant. New results are presented here to suggest that weight support through the feet and rapid extension of the legs are important proprioceptive inputs to initiate stepping. Our previous work has shown that infants can step at many different speeds when supported on a treadmill. The step cycle duration shortens as the speed increases, with the changes coming largely from the stance phase, just as in most other terrestrial animals. Moreover, we have shown that infants will step in all directions. Regardless of the direction of stepping, the step cycle changes in the same way with walking speed, suggesting the circuitry that controls different directions of walking share common elements. We have also shown that infant stepping is highly organized. Sensory inputs, whether proprioceptive or touch, are gated in a functional way so that only important sensory inputs generate a response. For example, touch to the lateral surface of the foot elicits a response only in sideways walking, and only in the leading limb. New data is presented here to show that the pattern generators from each limb can operate somewhat independently. On a split-belt treadmill with the 2 belts running at different speeds or in different directions, the legs showed considerable independence in behaviour. Yet, the pattern generators on each side interact to ensure that swing phase does not occur at the same time. These studies have provided insight into the organization of the pattern generator for walking in humans. It will be interesting in the future to study how maturation of the descending tracts changes walking behaviour to allow independent bipedal walking.     

Interlimb Coordination during Forward and Backward Walking in Primary School-Aged Children

Previous studies comparing forward (FW) and backward (BW) walking suggested that the leg kinematics in BW were essentially those of FW in reverse. This led to the proposition that in adults the neural control of FW and BW originates from the same basic neural circuitry. One aspect that has not received much attention is to what extent development plays a role in the maturation of neural control of gait in different directions. BW has been examined either in adults or infants younger than one year. Therefore, we questioned which changes occur in the intermediate phases (i.e. in primary school-aged children). Furthermore, previous research focused on the lower limbs, thereby raising the question whether upper limb kinematics are also simply reversed from FW to BW. Therefore, in the current study the emphasis was put both on upper and lower limb movements, and the coordination between the limbs. Total body 3D gait analysis was performed in primary school-aged children (N = 24, aged five to twelve years) at a preferred walking speed to record angular displacements of upper arm, lower arm, upper leg, lower leg, and foot with respect to the vertical (i.e. elevation angle). Kinematics and interlimb coordination were compared between FW and BW. Additionally, elevation angle traces of BW were reversed in time (revBW) and correlated to FW traces. Results showed that upper and lower limb kinematics of FW correlated highly to revBW kinematics in children, which appears to be consistent with the proposal that control of FW and BW may be similar. In addition, age was found to mildly alter lower limb kinematic patterns. In contrast, interlimb coordination was similar across all children, but was different compared to adults, measured for comparison. It is concluded that development plays a role in the fine-tuning of neural control of FW and BW.     

Inverted pendular running: a novel gait predicted by computer optimization is found between walk and run in birds

Idealized models of walking and running demonstrate that, energetically, walking should be favoured up to, and even somewhat over, those speeds and step lengths that can be achieved while keeping the stance leg under compression. Around these speeds, and especially with relatively long step lengths, computer optimization predicts a third, ‘hybrid’, gait: (inverted) pendular running (Srinivasan & Ruina 2006 Nature 439, 72–75 (doi:10.1038/nature04113)). This gait involves both walking-like vaulting mechanics and running-like ballistic paths. Trajectories of horizontal versus vertical centre of mass velocities—‘hodographs’—over the step cycle are distinctive for each gait: anticlockwise for walk; clockwise for run; figure-of-eight for the hybrid gait. Both pheasants and guineafowl demonstrate each gait at close to the predicted speed/step length combinations, although fully aerial ballistic phases are never achieved during the hybrid or ‘Grounded Inverted Pendular Running’ gait.     

Dynamic gait stability of treadmill versus overground walking in young adults

Treadmill has been broadly used in laboratory and rehabilitation settings for the purpose of facilitating human locomotion analysis and gait training. The objective of this study was to determine whether dynamic gait stability differs or resembles between the two walking conditions (overground vs. treadmill) among young adults. Fifty-four healthy young adults (age: 23.9 ± 4.7 years) participated in this study. Each participant completed five trials of overground walking followed by five trials of treadmill walking at a self-selected speed while their full body kinematics were gathered by a motion capture system. The spatiotemporal gait parameters and dynamic gait stability were compared between the two walking conditions. The results revealed that participants adopted a “cautious gait” on the treadmill compared with over ground in response to the possible inherent challenges to balance imposed by treadmill walking. The cautious gait, which was achieved by walking slower with a shorter step length, less backward leaning trunk, shortened single stance phase, prolonged double stance phase, and more flatfoot landing, ensures the comparable dynamic stability between the two walking conditions. This study could provide insightful information about dynamic gait stability control during treadmill ambulation in young adults.     

Torsion and Antero-Posterior Bending in the In Vivo Human Tibia Loading Regimes during Walking and Running

Bending, in addition to compression, is recognized to be a common loading pattern in long bones in animals. However, due to the technical difficulty of measuring bone deformation in humans, our current understanding of bone loading patterns in humans is very limited. In the present study, we hypothesized that bending and torsion are important loading regimes in the human tibia. In vivo tibia segment deformation in humans was assessed during walking and running utilizing a novel optical approach. Results suggest that the proximal tibia primarily bends to the posterior (bending angle: 0.15°–1.30°) and medial aspect (bending angle: 0.38°–0.90°) and that it twists externally (torsion angle: 0.67°–1.66°) in relation to the distal tibia during the stance phase of overground walking at a speed between 2.5 and 6.1 km/h. Peak posterior bending and peak torsion occurred during the first and second half of stance phase, respectively. The peak-to-peak antero-posterior (AP) bending angles increased linearly with vertical ground reaction force and speed. Similarly, peak-to-peak torsion angles increased with the vertical free moment in four of the five test subjects and with the speed in three of the test subjects. There was no correlation between peak-to-peak medio-lateral (ML) bending angles and ground reaction force or speed. On the treadmill, peak-to-peak AP bending angles increased with walking and running speed, but peak-to-peak torsion angles and peak-to-peak ML bending angles remained constant during walking. Peak-to-peak AP bending angle during treadmill running was speed-dependent and larger than that observed during walking. In contrast, peak-to-peak tibia torsion angle was smaller during treadmill running than during walking. To conclude, bending and torsion of substantial magnitude were observed in the human tibia during walking and running. A systematic distribution of peak amplitude was found during the first and second parts of the stance phase.     

Stability-normalised walking speed: A new approach for human gait perturbation research

In gait stability research, neither self-selected walking speeds, nor the same prescribed walking speed for all participants, guarantee equivalent gait stability among participants. Furthermore, these options may differentially affect the response to different gait perturbations, which is problematic when comparing groups with different capacities. We present a method for decreasing inter-individual differences in gait stability by adjusting walking speed to equivalent margins of stability (MoS). Eighteen healthy adults walked on a split-belt treadmill for two-minute bouts at 0.4 m/s up to 1.8 m/s in 0.2 m/s intervals. The stability-normalised walking speed (MoS = 0.05 m) was calculated using the mean MoS at touchdown of the final 10 steps of each speed. Participants then walked for three minutes at this speed and were subsequently exposed to a treadmill belt acceleration perturbation. A further 12 healthy adults were exposed to the same perturbation while walking at 1.3 m/s: the average of the previous group. Large ranges in MoS were observed during the prescribed speeds (6–10 cm across speeds) and walking speed significantly (P < 0.001) affected MoS. The stability-normalised walking speeds resulted in MoS equal or very close to the desired 0.05 m and reduced between-participant variability in MoS. The second group of participants walking at 1.3 m/s had greater inter-individual variation in MoS during both unperturbed and perturbed walking compared to 12 sex, height and leg length-matched participants from the stability-normalised walking speed group. The current method decreases inter-individual differences in gait stability which may benefit gait perturbation and stability research, in particular studies on populations with different locomotor capacities. [Preprint: https://doi.org/10.1101/314757]     

Wireless Tri-Axial Trunk Accelerometry Detects Deviations in Dynamic Center of Mass Motion Due to Running-Induced Fatigue

Small wireless trunk accelerometers have become a popular approach to unobtrusively quantify human locomotion and provide insights into both gait rehabilitation and sports performance. However, limited evidence exists as to which trunk accelerometry measures are suitable for the purpose of detecting movement compensations while running, and specifically in response to fatigue. The aim of this study was therefore to detect deviations in the dynamic center of mass (CoM) motion due to running-induced fatigue using tri-axial trunk accelerometry. Twenty runners aged 18–25 years completed an indoor treadmill running protocol to volitional exhaustion at speeds equivalent to their 3.2 km time trial performance. The following dependent measures were extracted from tri-axial trunk accelerations of 20 running steps before and after the treadmill fatigue protocol: the tri-axial ratio of acceleration root mean square (RMS) to the resultant vector RMS, step and stride regularity (autocorrelation procedure), and sample entropy. Running-induced fatigue increased mediolateral and anteroposterior ratios of acceleration RMS (p < .05), decreased the anteroposterior step regularity (p < .05), and increased the anteroposterior sample entropy (p < .05) of trunk accelerometry patterns. Our findings indicate that treadmill running-induced fatigue might reveal itself in a greater contribution of variability in horizontal plane trunk accelerations, with anteroposterior trunk accelerations that are less regular from step-to-step and are less predictable. It appears that trunk accelerometry parameters can be used to detect deviations in dynamic CoM motion induced by treadmill running fatigue, yet it is unknown how robust or generalizable these parameters are to outdoor running environments.     

The Combined Effects of Body Weight Support and Gait Speed on Gait Related Muscle Activity: A Comparison between Walking in the Lokomat Exoskeleton and Regular Treadmill Walking

Background

For the development of specialized training protocols for robot assisted gait training, it is important to understand how the use of exoskeletons alters locomotor task demands, and how the nature and magnitude of these changes depend on training parameters. Therefore, the present study assessed the combined effects of gait speed and body weight support (BWS) on muscle activity, and compared these between treadmill walking and walking in the Lokomat exoskeleton.

Methods

Ten healthy participants walked on a treadmill and in the Lokomat, with varying levels of BWS (0% and 50% of the participants’ body weight) and gait speed (0.8, 1.8, and 2.8 km/h), while temporal step characteristics and muscle activity from Erector Spinae, Gluteus Medius, Vastus Lateralis, Biceps Femoris, Gastrocnemius Medialis, and Tibialis Anterior muscles were recorded.

Results

The temporal structure of the stepping pattern was altered when participants walked in the Lokomat or when BWS was provided (i.e. the relative duration of the double support phase was reduced, and the single support phase prolonged), but these differences normalized as gait speed increased. Alternations in muscle activity were characterized by complex interactions between walking conditions and training parameters: Differences between treadmill walking and walking in the exoskeleton were most prominent at low gait speeds, and speed effects were attenuated when BWS was provided.

Conclusion

Walking in the Lokomat exoskeleton without movement guidance alters the temporal step regulation and the neuromuscular control of walking, although the nature and magnitude of these effects depend on complex interactions with gait speed and BWS. If normative neuromuscular control of gait is targeted during training, it is recommended that very low speeds and high levels of BWS should be avoided when possible.     

Validity of the MarkWiiR for kinematic analysis during walking and running gaits

The aim of this study was to validate the MarkWiiR (MW) captured by the Nintendo Wii-Remote (100-Hz) to assess active marker displacement by comparison with 2D video analysis. Ten participants were tested on a treadmill at different walking (1<6 km · h⁻¹) and running (10<13 km · h⁻¹) speeds. During the test, the active marker for MW and a passive marker for video analysis were recorded simultaneously with the two devices. The displacement of the marker on the two axes (x-y) was computed using two different programs, Kinovea 0.8.15 and CoreMeter, for the camera and MW, respectively. Pearson correlation was acceptable (x-axis r≥0.734 and y-axis r≥0.684), and Bland–Altman plots of the walking speeds showed an average error of 0.24±0.52% and 1.5±0.91% for the x- and y-axis, respectively. The difference of running speeds showed average errors of 0.67±0.33% and 1.26±0.33% for the x- and y-axes, respectively. These results demonstrate that the two measures are similar from both the x- and the y-axis perspective. In conclusion, these findings suggest that the MarkWiiR is a valid and reliable tool to assess the kinematics of an active marker during walking and running gaits.     

Effect of walking speed on inter-joint coordination differs between young and elderly adults

Investigating inter-joint coordination at different walking speeds in young and elderly adults could provide insights to age-related changes in neuromuscular control of gait. We examined effects of walking speed and age on the pattern and variability of inter-joint coordination. Gait analyses of 10 young and 10 elderly adults were performed with different self-selected speeds, including a preferred, faster, and slower speed. Continuous relative phase (CRP), derived from phase planes of two adjacent joints, was used to assess the inter-joint coordination. CRP patterns were examined with cross-correlation measures and root-mean-square (RMS) differences when comparing ensemble mean curves of the faster or slower speed to preferred speed walking. Variability of coordination for each participant was assessed with the average value of all standard deviations calculated for each data point over a gait cycle from all CRP curves, namely the deviation phase (DP). For hip-knee CRP pattern, RMS differences were significantly greater between the slower and preferred walking speeds than between the faster and preferred walking speeds in young adults, but this was not found in elderly adults. Significant group differences in RMS differences and cross-correlation measures were detected in hip-knee CRP patterns between the slower and preferred walking speeds. No significant walking speed or age effects were detected for the knee-ankle CRP. Significant walking speed effects were also detected in hip-knee DP values. However, no significant group differences were detected for all three speeds. These findings suggested that young and elder adults compromise changes of walking speed with different neuromuscular control strategies.     

Evidence for energy savings from aerial running in the Svalbard rock ptarmigan (Lagopus muta hyperborea)

Svalbard rock ptarmigans were walked and run upon a treadmill and their energy expenditure measured using respirometry. The ptarmigan used three different gaits: a walking gait at slow speeds (less than or equal to 0.75 m s(-1)), grounded running at intermediate speeds (0.75 m s(-1) < U < 1.67 m s(-1)) and aerial running at high speeds (greater than or equal to 1.67 m s(-1)). Changes of gait were associated with reductions in the gross cost of transport (COT; J kg(-1) m(-1)), providing the first evidence for energy savings with gait change in a small crouched-postured vertebrate. In addition, for the first time (excluding humans) a decrease in absolute metabolic energy expenditure (rate of O(2) consumption) in aerial running when compared with grounded running was identified. The COT versus U curve varies between species and the COT was cheaper during aerial running than grounded running, posing the question of why grounded running should be used at all. Existing explanations (e.g. stability during running over rocky terrain) amount to just so stories with no current evidence to support them. It may be that grounded running is just an artefact of treadmill studies. Research investigating the speeds used by animals in the field is sorely needed.     

Gait selection in the ostrich: mechanical and metabolic characteristics of walking and running with and without an aerial phase

It has been argued that minimization of metabolic-energy costs is a primary determinant of gait selection in terrestrial animals. This view is based predominantly on data from humans and horses, which have been shown to choose the most economical gait (walking, running, galloping) for any given speed. It is not certain whether a minimization of metabolic costs is associated with the selection of other prevalent forms of terrestrial gaits, such as grounded running (a widespread gait in birds). Using biomechanical and metabolic measurements of four ostriches moving on a treadmill over a range of speeds from 0.8 to 6.7 m s(-1), we reveal here that the selection of walking or grounded running at intermediate speeds also favours a reduction in the metabolic cost of locomotion. This gait transition is characterized by a shift in locomotor kinetics from an inverted-pendulum gait to a bouncing gait that lacks an aerial phase. By contrast, when the ostrich adopts an aerial-running gait at faster speeds, there are no abrupt transitions in mechanical parameters or in the metabolic cost of locomotion. These data suggest a continuum between grounded and aerial running, indicating that they belong to the same locomotor paradigm.     

Noisy Interlimb Coordination Can Be a Main Cause of Freezing of Gait in Patients with Little to No Parkinsonism

Freezing of gait in patients with Parkinson’s disease is associated with several factors, including interlimb incoordination and impaired gait cycle regulation. Gait analysis in patients with Parkinson’s disease is confounded by parkinsonian symptoms such as rigidity. To understand the mechanisms underlying freezing of gait, we compared gait patterns during straight walking between 9 patients with freezing of gait but little to no parkinsonism (freezing patients) and 11 patients with Parkinson’s disease (non-freezing patients). Wireless sensors were used to detect foot contact and toe-off events, and the step phase of each foot contact was calculated by defining one stride cycle of the other leg as 360°. Phase-resetting analysis was performed, whereby the relation between the step phase of one leg and the subsequent phase change in the following step of the other leg was quantified using regression analysis. A small slope of the regression line indicates a forceful correction (phase reset) at every step of the deviation of step phase from the equilibrium phase, usually at around 180°. The slope of this relation was smaller in freezing patients than in non-freezing patients, but the slope exhibited larger step-to-step variability. This indicates that freezing patients executed a forceful but noisy correction of the deviation of step phase, whereas non-freezing patients made a gradual correction of the deviation. Moreover, freezing patients tended to show more variable step phase and stride time than non-freezing patients. Dynamics of a model of two coupled oscillators interacting through a phase resetting mechanism were examined, and indicated that the deterioration of phase reset by noise provoked variability in step phase and stride time. That is, interlimb coordination can affect regulation of the gait cycle. These results suggest that noisy interlimb coordination, which probably caused forceful corrections of step phase deviation, can be a cause of freezing of gait.     

Effects of walking speed, strength and range of motion on gait stability in healthy older adults

Falls pose a tremendous risk to those over 65 and most falls occur during locomotion. Older adults commonly walk slower, which many believe helps improve walking stability. While increased gait variability predicts future fall risk, increased variability is also caused by walking slower. Thus, we need to better understand how differences in age and walking speed independently affect dynamic stability during walking. We investigated if older adults improved their dynamic stability by walking slower, and how leg strength and flexibility (passive range of motion (ROM)) affected this relationship. Eighteen active healthy older and 17 healthy younger adults walked on a treadmill for 5min each at each of 5 speeds (80-120% of preferred). Local divergence exponents and maximum Floquet multipliers (FM) were calculated to quantify each subject's inherent local dynamic stability. The older subjects walked with the same preferred walking speeds as the younger subjects (p=0.860). However, these older adults still exhibited greater local divergence exponents (p<0.0001) and higher maximum FM (p<0.007) than the younger adults at all walking speeds. These older adults remained more locally unstable (p<0.04) even after adjusting for declines in both strength and ROM. In both age groups, local divergence exponents decreased at slower speeds and increased at faster speeds (p<0.0001). Maximum FM showed similar changes with speed (p<0.02). Both younger and older adults exhibited decreased instability by walking slower, in spite of increased variability. These increases in dynamic instability might be more sensitive indicators of future fall risk than changes in gait variability.     

Synergistic Structure in the Speed Dependent Modulation of Muscle Activity in Human Walking

Recently, a modular organisation has been proposed to simplify control of the large number of muscles involved in human walking. Although previous research indicates that a single set of modular activation patterns can account for muscle activity at different speeds, these studies only provide indirect evidence for the idea that speed regulation in human walking is under modular control. Here, a more direct approach was taken to assess the synergistic structure that underlies speed regulation, by isolating speed effects through the construction of gain functions that represent the linear relation between speed and amplitude for each point in the time-normalized gait cycle. The activity of 13 muscles in 13 participants was measured at 4 speeds (0.69, 1.00, 1.31, and 1.61 ms^-1) during treadmill walking. Gain functions were constructed for each of the muscles, and gain functions and the activity patterns at 1.00 ms^-1 were both subjected to dimensionality reduction, to obtain modular gain functions and modular basis functions, respectively. The results showed that 4 components captured most of the variance in the gain functions (74.0% ± 1.3%), suggesting that the neuromuscular regulation of speed is under modular control. Correlations between modular gain functions and modular basis functions (range 0.58–0.89) and the associated synergistic muscle weightings (range 0.6–0.95) were generally high, suggesting substantial overlap in the synergistic control of the basic phasing of muscle activity and its modulation through speed. Finally, the combined set of modular functions and associated weightings were well capable of predicting muscle activity patterns obtained at a speed (1.31 ms^-1) that was not involved in the initial dimensionality reduction, confirming the robustness of the presently used approach. Taken together, these findings provide direct evidence of synergistic structure in speed regulation, and may inspire further work on flexibility in the modular control of gait.     

The influence of body weight support on ankle mechanics during treadmill walking

The use of body weight support (BWS) systems during locomotor retraining has become routine in clinical settings. BWS alters load receptor feedback, however, and may alter the biomechanical role of the ankle plantarflexors, influencing gait. The purpose of this study was to characterize the biomechanical adaptations that occur as a result of a change in limb load (controlled indirectly through BWS) and gait speed during treadmill locomotion. Fifteen unimpaired participants underwent gait analysis with surface electromyography while walking on an instrumented dual-belt treadmill at seven different speeds (ranging from 0.4 to 1.6 m/s) and three BWS conditions (ranging from 0% to 40% BWS). While walking, spatiotemporal measures, anterior/posterior ground reaction forces, and ankle kinetics and muscle activity were measured and compared between conditions. At slower gait speeds, propulsive forces and ankle kinetics were unaffected by changing BWS; however, at gait speeds ≥approximately 0.8 m/s, an increase in BWS yielded reduced propulsive forces and diminished ankle plantarflexor moments and powers. Muscle activity remained unaltered by changing BWS across all gait speeds. The use of BWS could provide the advantage of faster walking speeds with the same push-off forces as required of a slower speed. While the use of BWS at slower speeds does not appear to detrimentally affect gait, it may be important to reduce BWS as participants progress with training, to encourage maximal push-off forces. The reduction in plantarflexor kinetics at higher speeds suggests that the use of BWS in higher functioning individuals may impair the ability to relearn walking.