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Bruijn, S. M.

Publications and source records attributed to Bruijn, S. M..

At least 37 records · Page 2Linked to original sources

The energetic effect of hip flexion and retraction in walking at different speeds: a modeling study

In human walking, power for propulsion is generated primarily via ankle and hip muscles. The addition of a passive hip spring to simple bipedal models appears more efficient than using only push-off impulse, at least, when hip spring associated energetic costs are not considered. Hip flexion and retraction torques, however, are not free, as they are produced by muscles demanding metabolic energy. Studies evaluating the inclusion of hip actuation costs, especially during the swing phase, and the hip actuations energetic benefits are few and far between. It is also unknown whether these possible benefits/effects may depend on speed. We simulated a planar flat-feet model walking stably over a range of speeds. We asked whether the addition of independent hip flexion and retraction remains energetically beneficial when considering work-based metabolic cost of transport (MCOT) with different efficiencies of doing positive and negative work. We found asymmetric hip actuation can reduce the estimated MCOT relative to ankle actuation by up to 6%, but only at medium speeds. The corresponding optimal strategy is zero hip flexion and some hip retraction actuation. The reason for this reduced MCOT is a reduction in collision loss being larger than the associated increase in hip negative work. Both terms require positive mechanical work to compensate, yielding a larger reduction in metabolic work per step than the reduction in step length. Our study shows how ankle actuation, hip flexion, and retraction actuation can be coordinated to reduce MCOT.

physiology↗

Differential effects of ankle constraints on foot placement control between normal and split belt treadmills

Mediolateral ankle moment control contributes to gait stability. Ankle moments can be constrained by walking with a shoe with a ridge underneath the sole, narrowing the mediolateral support surface. In our previous study, such ankle moment constraints resulted in an increased step width and a decrease in the degree of foot placement control, as defined by the percentage of variance in foot placement that can be explained by CoM state. However, since our previous study was performed on a split-belt treadmill and the narrow ridge could fit inside the gap between the belts, it is not evident whether these effects can be attributed to the constrained ankle moment control or to avoidance of this gap. Therefore, we investigated if the effects of ankle moment constraints are dependent on whether participants walk on a normal treadmill or a split-belt treadmill. We included fourteen healthy young adults. Walking with constrained ankle moment control resulted in a wider step width on both treadmills. Yet, the increase in step width was larger on the split-belt treadmill compared to on the normal treadmill. We only found a decreased degree of foot placement control on the split-belt treadmill, whilst the degree of foot placement control increased on the normal treadmill. We conclude that the effects of ankle moment constraints reported in our previous study were confounded by the use of a split-belt treadmill. For future research, we recommend using a normal treadmill whenever possible, because the gap in a split-belt treadmill might affect gait parameters.

biophysics↗

The effect of external lateral stabilization on ankle moment control during steady-state walking

External lateral stabilization can help identify stability control mechanisms during steady-state walking. The degree of step-by-step foot placement control and step width are known to decrease when walking with external lateral stabilization. Here, we investigated the effect of external lateral stabilization on ankle moment control in healthy participants. Ankle moment control complements foot placement, by allowing a corrective center-of-pressure shift once the foot has been placed. This is reflected by a model predicting this center-of-pressure shift based on the preceding foot placement error. Here, the absolute explained variance accounted for by this model decreased when walking with external lateral stabilization. In other words, we found a reduction in the contribution of step-by-step ankle moment control to mediolateral gait stability when externally stabilized. Concurrently, foot placement error and the average center-of-pressure shift remained unchanged.

biophysics↗

Does ankle push-off correct for errors in anterior-posterior foot placement relative to center-of-mass state?

I.Understanding the mechanisms humans use to stabilize walking is vital for predicting falls in elderly. Modeling studies identified two potential mechanisms to stabilize gait in the anterior-posterior direction: foot placement control and ankle push-off control: Foot placement depends on position and velocity of the center-of-mass (CoM) and push-off modulates with deviations between actual and predicted CoM trajectories. While both control mechanisms have been reported in humans, it is unknown whether especially the latter one is employed in unperturbed steady-state walking. On the one hand, the covariance between CoM states and anterior-posterior foot placement served as a measure of foot placement control. On the other hand, we determined variations in ankle push-off as a function of differences between the actual foot placement and the one predicted from CoM states. We estimated the corresponding correlations and consider them indictors for push-off control based on foot placement errors. We found ankle push-off torque to be correlated to the foot placement error in 30 participants when walking at normal and slow speed, with mean correlation values of up to 0.45. Our study suggests that humans use a push-off strategy for correcting foot placement errors in steady-state walking.

physiology↗

Limited effects of age on the use of the ankle and counter-rotation mechanism in the sagittal plane

Two mechanisms can be used to accelerate the center of mass (CoM) to control the CoM in relation to the base of support during standing. The first is applying ankle moments to shift the center of pressure (CoP), which has been coined the "CoP mechanism". The second is changing the angular momentum around the CoM to change the direction of the ground reaction force, i.e., the "counter-rotation mechanism". At both the beginning and the end of the lifespan, problems with postural control are common. In this study, we asessed anteroposterior balance performance and the related use of these postural control mechanisms in children, younger adults, and older adults. Sixteen pre-pubertal children (6-9y), 17 younger adults (18-24y) and eight older adults (65-80y) performed bipedal upright standing trials of 16 seconds on a rigid surface and on three balance boards that could freely move in the sagittal plane, varying in height (15-19 cm) of the board above the point of contact with the floor. Full body kinematics were measured. Performance related outcome measures, i.e., the number of trials with balance loss and the Root Mean Square (RMS) of the time series of the CoM acceleration were calculated. Additionally, the RMS of the time series of the CoM acceleration due to the CoP and counter-rotation mechanism and the contributions of the CoP and the counter-rotation mechanism to the CoM acceleration (in %) in the sagittal plane were calculated. Furthermore, selected kinematic measures, i.e., the orientation of the board and the head and the Mean Power Frequency of balance board orientation and of CoM acceleration were calculated. Compared to younger adults, children and older adults showed a poorer balance performance, reflected by a greater RMS of CoM accelerations and more balance loss in older adults. Across age groups and conditions, the contribution of the CoP mechanism to the total CoM acceleration was dominant, i.e., 95%-108%. The contribution of the counter-rotation mechanism ranged between 19%-31% (with totals higher than 100% indicating opposite effects of both mechanisms). We suggest that the contribution of the counter-rotation mechanism is limited, since the counter-rotation mechanism would conflict with stabilizing the orientation of the head in space. Furthermore, children did use the counter-rotation mechanism relatively more to accelerate the CoM compared to younger adults. Possibly this reflects that they are still learning to limit the contribution of the counter-rotation mechanism to the same extent as adults.

bioengineering↗

The effect of constraining mediolateral ankle moments and foot placement on the use of the counter-rotation mechanism during walking

Stability during walking can be maintained by shifts of the Center of Pressure through modulation of foot placement and ankle moments (CoP-mechanism). An additional mechanism to stabilize gait, is the counter-rotation mechanism i.e. changing the angular momentum of segments around the Center of Mass (CoM) to change the direction of the ground reaction force. It is unknown if and how humans use the counter-rotation mechanism to control the CoM during walking and how this interacts with the CoP-mechanism. Thirteen healthy adults walked on a treadmill, while full-body kinematic and force plate data were obtained. The contributions of the CoP and the counter-rotation mechanisms to control the CoM were calculated during steady-state walking, walking on LesSchuh, i.e. constraining mediolateral CoP shifts underneath the stance foot and walking on LesSchuh at 50% of normal step width, constraining both foot placement and ankle mechanisms (LesSchuh50%). A decreased magnitude of within-stride control by the CoP-mechanism was compensated for by an increased magnitude of within-stride control by the counter-rotation mechanism during LesSchuh50% compared to steady-state walking. This suggests that the counter-rotation mechanism is used to stabilize gait when needed. However, the mean contribution of the counter-rotation mechanism over strides did not increase during LesSchuh50% compared to steady-state walking. The CoP-mechanism was the main contributor to the total CoM acceleration. The use of the counter-rotation mechanism may be limited because angular accelerations ultimately need to be reversed and because of interference with other task constraints, such as head stabilization and preventing interference with the gait pattern.

bioengineering↗

Effects of vestibular stimulation on gait stability and variability

Vestibular information modulates muscle activity during gait, presumably to contribute to stability. If this is the case, stronger effects of perturbing vestibular information on local dynamic stability of gait, a measure of the locomotor systems response to small, naturally occurring perturbations, can be expected for narrow-base walking (which needs more control) than for normal walking and smaller effects for wide-base walking (which needs less control). An important mechanism to stabilize gait is to coordinate foot placement to center of mass (CoM) state. Vestibular information most likely contributes to sensing this CoM state. We, therefore, expected that stochastic electrical vestibular stimulation (EVS) would decrease the correlation between foot placement and CoM state during the preceding swing phase. In fourteen healthy participants, we measured the kinematics of the trunk (as a proxy of the CoM), and feet, while they walked on a treadmill in six conditions: control (usual step width), narrow-base, and wide-base, each with and without stochastic EVS (peak amplitude of 5 mA; RMS of ~ 1.2 mA; frequency band from 0 to 25 Hz). Stochastic EVS decreased local dynamic stability irrespective of step width. Foot placement correlated stronger with trunk motion during walking with EVS than without in the control condition. However, residual variance in foot placement was increased when walking with EVS, indicating less precise foot placement. Thus, a vestibular error signal leads to a decrease in gait stability and precision of foot placement but these effects are not consistently modulated by step width.

neuroscience↗

Effects of age and surface instability on the control of the center of mass

During standing, posture can be controlled by accelerating the Center of Mass (CoM) through shifting the center of pressure (CoP) within the base of support by applying ankle moments ("CoP mechanism"), or through the "counter-rotation mechanism", i.e., changing the angular momentum of segments around the CoM to change the direction of the ground reaction force. Postural control develops over the lifespan; at both the beginning and the end of the lifespan adequate postural control appears more challenging. In this study, we aimed to assess mediolateral balance performance and the related use of the postural control mechanisms in children, older adults and young adults when standing on different (unstable) surfaces. Sixteen pre-pubertal children (6-9y), 17 young adults (18-24y) and eight older adults (65-80y) performed bipedal upright standing trials of 16 seconds on a rigid surface and on three balance boards that could freely move in the frontal plane, varying in height (15-19 cm) of the surface of the board above the point of contact with the floor. Full body kinematics (16 segments, 48 markers, using SIMI 3D-motion analysis system (GmbH) and DeepLabCut and Anipose) were retrieved. Performance related outcome measures, i.e., the number of trials with balance loss and the Root Mean Square (RMS) of the time series of the CoM acceleration, the contributions of the CoP mechanism and the counter-rotation mechanism to the CoM acceleration in the frontal plane and selected kinematic measures, i.e. the orientation of the board and the head and the Mean Power Frequency (MPF) of the balance board orientation and the CoM acceleration were determined. Balance loss only occurred when standing on the highest balance board, twice in one older adult once in one young adult. In children and older adults, the RMS of the CoM accelerations were larger, corresponding to poorer balance performance. Across age groups and conditions, the contribution of the CoP mechanism to the total CoM acceleration was much larger than that of the counter-rotation mechanisms, ranging from 94%-113% vs 23%-38% (with totals higher than 100% indicating opposite effects of both mechanisms). Deviations in head orientation were small compared to deviations in balance board orientation. We hypothesize that the CoP mechanism is dominant, since the counter-rotation mechanism would conflict with stabilizing the orientation of the head in space.

bioengineering↗

Can foot placement during gait be trained? Adaptations in stability control when ankle moments are constrained

Accurate coordination of mediolateral foot placement, relative to the center of mass kinematic state, is one of the mechanisms which ensures mediolateral stability during human walking. Previously, we found that shoes constraining ankle moments decreased the degree of foot placement control with respect to the center of mass kinematic state. As such, ankle moment constraints can be seen as a perturbation of foot placement. Direct mechanical perturbations of the swing leg trajectory can improve the degree of foot placement control as an after-effect. Here, we asked whether constrained ankle moments could have a similar effect. If confirmed, this would offer a simple training tool for individuals with impaired foot placement control. Participants walked in three conditions; normal (baseline) while wearing shoes constraining ankle moments (training) and normal again (after-effects). The degree of foot placement control was calculated as the percentage of variance in foot placement that could be predicted based on the center of mass kinematic state in the preceding swing phase. During training, the degree of foot placement control decreased initially compared to baseline, but it gradually improved over time. In the after-effect condition, it was higher than during baseline, yet not significantly so. During training, we observed increased step width, decreased stride time and reduced local dynamic stability. In conclusion, constraining ankle moment control deteriorates the degree of foot placement control. A non-significant trend towards an improved degree of foot placement control after prolonged exposure to constrained ankle moments, allows for speculation on a training potential.

biophysics↗

Contribution of arm movements to recovery after a trip in older adults.

Falls are common in daily life, often caused by trips and slips and, particularly in older adults, with serious consequences. Although arm movements play an important role in balance control, there is limited research into the role of arm movements during balance recovery after tripping in older adults. We investigated how older adults use their arms to recover from a trip and the difference in the effects of arm movements between fallers (n=5) and non-fallers (n=11). Sixteen older males and females (69.7{+/-}2.3 years) walked along a walkway and were occasionally tripped over suddenly appearing obstacles. We analysed the first trip using a biomechanical model based on full-body kinematics and force-plate data to calculate whole body orientation during the trip and recovery phase. With this model, we simulated the effects of arm movements at foot-obstacle impact and during trip recovery on body orientation. Apart from an increase in sagittal plane forward body rotation at touchdown in fallers, we found no significant differences between fallers and non-fallers in the effects of arm movements on trip recovery. Like earlier studies in young adults, we found that arm movements during the recovery phase had most favourable effects in the transverse plane: by delaying the transfer of angular momentum of the arms to the body, older adults rotated the tripped side more forward thereby allowing for a larger recovery step. Older adults that are prone to falling might improve their balance recovery after tripping by learning to prolong ongoing arm movements.

animal behavior and cognition↗

A new protocol for multiple muscle mapping using nTMS

The representation of muscles in the cortex can be mapped using navigated transcranial magnetic stimulation. The mapping can be quantified via measures like the centre of gravity or the size of the region of excitability. Determining these measures typically relies only on stimulation points that yield motor-evoked potentials (MEPs); stimulations that do not elicit an MEP, i.e., non-MEP points, are ignored entirely. In this study, we show how incorporating non-MEP points may affect the estimates of the size and centroid of the excitable area in eight hand and forearm muscles after mono-phasic single-pulse TMS. We performed test-retest assessments in twenty participants and estimated the reliability of centroids and sizes of the corresponding areas using inter-class correlation coefficients. For most muscles, the reliability turned out good, if not excellent. As expected, removing the non-MEP points significantly decreased area sizes and area weights, suggesting that conventional approaches that do not account for non-MEP points are likely to overestimate the regions of excitability.

neuroscience↗

Balance training improves feedback control of perturbed balance in older adults

Recovering balance after perturbations becomes challenging with aging, but an effective balance training could reduce such challenges. In this study, we examined the effect of balance training on feedback control after unpredictable perturbations by investigating balance performance, recovery strategy, and muscle synergies. We assessed the effect of balance training on unipedal perturbed balance in twenty older adults (>65 years) after short-term (one session) and long-term (3-weeks) training. Participants were exposed to random medial and lateral perturbations consisting of 8-degree rotations of a robot-controlled balance platform. We measured full-body 3D kinematics and activation of 9 muscles (8 stance leg muscles, one trunk muscle) during 2.5 s after the onset of perturbation. The perturbation was divided into 3 phases: phase1 from the onset to maximum rotation of the platform, phase 2 from the maximum rotation angle to the 0-degree angle and phase 3 after platform movement. Balance performance improved after long-term training as evidenced by decreased amplitudes of center of mass acceleration and rate of change of body angular momentum. The rate of change of angular momentum did not directly contribute to return of the center of mass within the base of support, but it reoriented the body to an aligned and vertical position. The improved performance coincided with altered activation of synergies depending on the direction and phase of the perturbation. We concluded that balance training improves control of perturbed balance, and reorganizes feedback responses, by changing temporal patterns of muscle activation. These effects were more pronounced after long-term than short-term training.

neuroscience↗

Ankle muscles activate: mediolateral center of pressure control ensures stable gait

During steady-state walking, mediolateral gait stability can be maintained by controlling the center of pressure (CoP). The CoP modulates the moment of the ground reaction force, which brakes and reverses movement of the center of mass (CoM) towards the lateral border of the base of support. In addition to foot placement, ankle moments serve to control the CoP. We hypothesized that, during steady-state walking, single stance ankle moments establish a CoP shift to correct for errors in foot placement. We expected ankle muscle activity to be associated with this complementary CoP shift. During treadmill walking, full-body kinematics, ground reaction forces and electromyography were recorded in thirty healthy participants. We found a negative relationship between preceding foot placement error and CoP displacement during single stance; steps that were too medial were compensated for by a lateral CoP shift and vice versa, steps that were too lateral were compensated for by a medial CoP shift. Peroneus longus, soleus and tibialis anterior activity correlated with these CoP shifts. As such, we identified an (active) ankle strategy during steady-state walking. As expected, absolute explained CoP variance by foot placement error decreased when walking with shoes constraining ankle moments. Yet, contrary to our expectations that ankle moment control would compensate for constrained foot placement, the absolute explained CoP variance by foot placement error did not increase when foot placement was constrained. We argue that this lack of compensation reflects the interdependent nature of ankle moment and foot placement control. We suggest that single stance ankle moments do not only compensate for preceding foot placement errors, but also assist control of the subsequent foot placement. Foot placement and ankle moment control are caught in a circular relationship, in which constraints imposed on one will also influence the other.

neuroscience↗

Differences in gait stability, trunk and foot accelerations between healthy young and older women

Our aim was to evaluate differences in gait acceleration intensity, variability and stability of feet and trunk between older females and young females using inertial sensors. Twenty older females (OF; mean age 68.4, SD 4.1 years) and eighteen young females (YF; mean age 22.3, SD 1.7 years) were asked to walk straight for 100 meters at their preferred speed, while wearing inertial sensors on heels and lower back. We calculated spatiotemporal measures, foot and trunk acceleration characteristics and their variability, as well as trunk stability using the local divergence exponent (LDE). Two-way analysis of variance (including the factors foot and age), Students t-test, and Mann-Whitney U test were used to compare statistical differences of measures between groups. Cohens d effects were calculated for each variable. Foot maximum vertical acceleration and amplitude, trunk-foot vertical acceleration attenuation, as well as their variability were significantly smaller in OF than in YF. In contrast, trunk mediolateral acceleration amplitude, maximum vertical acceleration, and amplitude, as well as their variability were significantly larger in OF than in YF. Moreover, OF showed lower stability (i.e., higher LDE values) in mediolateral acceleration, mediolateral and vertical angular velocity of the trunk. Even though we measured healthy older females, these participants showed lower vertical foot accelerations with higher vertical trunk acceleration, lower trunk-foot vertical acceleration attenuation, less gait stability, and more variability of the trunk, and hence, were more likely to fall. These findings suggest that instrumented gait measurements may help for early detection of changes or impairments in gait performance, even before this can be observed by clinical eye or gait speed.

bioengineering↗

Neuromuscular control of gait stability in older adults is adapted to environmental demands but not improved after standing balance training

Balance training aims to improve balance and transfer acquired skills to real-life tasks and conditions. How older adults adapt gait control to different conditions, and whether these adaptations are altered by balance training remains unclear. We investigated adaptations in neuromuscular control of gait in twenty-two older adults (72.6 {+/-} 4.2 years) between normal (NW) and narrow-base walking (NBW), and the effects of a standing balance training program shown to enhance unipedal balance control in the same participants. At baseline, after one session and after 3-weeks of training, kinematics and EMG of NW and NBW on a treadmill were measured. Gait parameters and temporal activation profiles of five synergies extracted from 11 muscles were compared between time-points and gait conditions. No effects of balance training or interactions between training and walking condition on gait parameters or synergies were found. Trunk center of mass (CoM) displacement and velocity (vCoM), and the local divergence exponent (LDE), were lower in NBW compared to NW. For synergies associated with stance of the non-dominant leg and weight acceptance of the dominant leg, full width at half maximum (FWHM) of the activation profiles was smaller in NBW compared to NW. For the synergy associated with non-dominant heel strike, FWHM was greater in NBW compared to NW. The Center of Activation (CoA) of the activation profile associated with dominant leg stance occurred earlier in NBW compared to NW. CoAs of activation profile associated with non-dominant stance and non-dominant and dominant heel strikes were delayed in NBW compared to NW. The adaptations of synergies to NBW can be interpreted as related to a more cautious weight transfer to the new stance leg and enhanced control over CoM movement in the stance phase. However, control of mediolateral gait stability and these adaptations were not affected by balance training.

neuroscience↗

Stabilization demands of walking modulate the vestibular contributions to gait

Stable walking relies critically on motor responses to signals of head motion provided by the vestibular system, which are phase-dependent and modulated differently within each muscle. It is unclear, however, whether these vestibular contributions also vary according to the stability of the walking task. Here we investigate how vestibular signals influence muscles relevant for gait stability (medial gastrocnemius, gluteus medius and erector spinae) - as well as their net effect on ground reaction forces - while humans walked normally, with mediolateral stabilization, wide and narrow steps. We estimated local dynamic stability of trunk kinematics together with coherence of electrical vestibular stimulation (EVS) with muscle activity and mediolateral ground reaction forces. Walking with external stabilization increased local dynamic stability and decreased coherence between EVS and all muscles/forces compared to normal walking. Wide-base walking also decreased vestibulomotor coherence, though local dynamic stability did not differ. Conversely, narrow-base walking increased local dynamic stability, but produced muscle-specific increases and decreases in coherence that resulted in a net increase in vestibulomotor coherence with ground reaction forces. Overall, our results show that while vestibular contributions may vary with gait stability, they more critically depend on the stabilization demands (i.e. control effort) needed to maintain a stable walking pattern.

neuroscience↗

The underlying mechanisms of improved balance aftershort- and long-term training in older adults

Training improves balance control in older adults, but the time course and neural mechanisms underlying these improvements are unclear. We studied balance robustness and performance, H-reflex gains, paired reflex depression, and co-contraction duration in ankle muscles after one and ten training sessions in 22 older adults (+65yrs). Mediolateral balance robustness, time to balance loss in unipedal standing on a platform with decreasing rotational stiffness, improved (33%) after one session, with no further improvement after ten sessions. Balance performance, absolute mediolateral center of mass velocity, improved (18.75%) after one session in perturbed unipedal standing and (18.18%) after ten sessions in unperturbed unipedal standing. Co-contraction duration of soleus/tibialis anterior increased (16%) after ten sessions. H-reflex gain and paired reflex depression excitability did not change. H-reflex gains were lower, and soleus/tibialis anterior co-contraction duration was higher in participants with more robust balance after ten sessions, and co-contraction duration was higher in participants with better balance performance at several time-points. Changes in robustness and performance were uncorrelated with changes in co-contraction duration, H-reflex gain, or paired reflex depression. In older adults, balance robustness improved over a single session, while performance improved gradually over multiple sessions. Changes in co-contraction and excitability of ankle muscles were not exclusive causes of improved balance. HighlightsO_LIBalance robustness and balance performance in perturbed unipedal standing was improved after one balance training session, with no further improvement after ten sessions. C_LIO_LIBalance performance in unperturbed unipedal standing was improved after ten sessions C_LIO_LIH-reflex and paired reflex depression did not change after training in unipedal or bipedal standing. C_LIO_LICo-contraction duration of antagonistic ankle muscles increased after ten sessions in perturbed and unperturbed unipedal standing. C_LIO_LIChanges in co-contraction duration and excitability of ankle muscles were not exclusive causes of improved balance. C_LI

neuroscience↗

Step width and frequency to modulate: Active foot placement control ensures stable gait

Step-by-step foot placement control, relative to the center of mass (CoM) kinematic state, is generally considered a dominant mechanism for maintenance of gait stability. By adequate (mediolateral) positioning of the center of pressure with respect to the CoM, the ground reaction force generates a moment that prevents falling. In healthy individuals, foot placement is complemented mainly by ankle moment control ensuring stability. To evaluate possible compensatory relationships between step-by-step foot placement and complementary ankle moments, we investigated the degree of (active) foot placement control during steady-state walking, and under either foot placement-, or ankle moment constraints. Thirty healthy participants walked on a treadmill, while full-body kinematics, ground reaction forces and EMG activities were recorded. As a replication of earlier findings, we first showed step-by-step foot placement is associated with preceding CoM state and hip ab-/adductor activity during steady-state walking. Tight control of foot placement appears to be important at normal walking speed because there was a limited change in the degree of foot placement control despite the presence of a foot placement constraint. At slow speed, the degree of foot placement control decreased substantially, suggesting that tight control of foot placement is less essential when walking slowly. Step-by-step foot placement control was not tightened to compensate for constrained ankle moments. Instead compensation was achieved through increases in step width and stride frequency.

neuroscience↗