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.     

Correlations of pelvis state to foot placement do not imply within-step active control

Experimental studies of human walking have shown that within an individual step, variations in the center of mass (CoM) state can predict corresponding variations in the next foot placement. This has been interpreted by some to indicate the existence of active control in which the nervous system uses the CoM state at or near mid-stance to regulate subsequent foot placement. However, the passive dynamics of the moving body and/or moving limbs also contribute (perhaps strongly) to foot placement, and thus to its variation. The extent to which correlations of CoM state to foot placement reflect the effects of within-step active control, those of passive dynamics, or some combination of both, remains an important and still open question. Here, we used an open-loop-stable 2D walking model to show that this predictive ability cannot by itself be taken as evidence of within-step active control. In our simulations, we too find high correlations between the CoM state and subsequent foot placement, but these correlations are entirely due to passive dynamics as our system has no active control, either within a step or between steps. This demonstrates that any inferences made from such correlations about within-step active control require additional supporting evidence beyond the correlations themselves. Thus, these within-step predictive correlations leave unresolved the relative importance of within-step active control as compared to passive dynamics, meaning that such methods should be used to characterize control in human walking only with caution.     

Running backwards: soft landing–hard takeoff,a less efficient rebound

Human running at low and intermediate speeds is characterized by a greater average force exerted after ‘landing’, when muscle–tendon units are stretched (‘hard landing’), and a lower average force exerted before ‘takeoff’, when muscle–tendon units shorten (‘soft takeoff’). This landing–takeoff asymmetry is consistent with the force–velocity relation of the ‘motor’ (i.e. with the basic property of muscle to resist stretching with a force greater than that developed during shortening), but it may also be due to the ‘machine’ (e.g. to the asymmetric lever system of the foot operating during stance). Hard landing and soft takeoff—never the reverse—were found in running, hopping and trotting animals using diverse lever systems, suggesting that the different machines evolved to comply with the basic force–velocity relation of the motor. Here we measure the mechanical energy of the centre of mass of the body in backward running, an exercise where the normal coupling between motor and machine is voluntarily disrupted, in order to see the relevance of the motor–machine interplay in human running. We find that the landing–takeoff asymmetry is reversed. The resulting ‘soft landing’ and ‘hard takeoff’ are associated with a reduced efficiency of positive work production. We conclude that the landing–takeoff asymmetry found in running, hopping and trotting is the expression of a convenient interplay between motor and machine. More metabolic energy must be spent in the opposite case when muscle is forced to work against its basic property (i.e. when it must exert a greater force during shortening and a lower force during stretching).     

The bounce of the body in hopping,running and trotting: different machines with the same motor

The bouncing mechanism of human running is characterized by a shorter duration of the brake after ‘landing’ compared with a longer duration of the push before ‘takeoff’. This landing–takeoff asymmetry has been thought to be a consequence of the force–velocity relation of the muscle, resulting in a greater force exerted during stretching after landing and a lower force developed during shortening before takeoff. However, the asymmetric lever system of the human foot during stance may also be the cause. Here, we measure the landing–takeoff asymmetry in bouncing steps of running, hopping and trotting animals using diverse lever systems. We find that the duration of the push exceeds that of the brake in all the animals, indicating that the different lever systems comply with the basic property of muscle to resist stretching with a force greater than that developed during shortening. In addition, results show both the landing–takeoff asymmetry and the mass-specific vertical stiffness to be greater in small animals than in large animals. We suggest that the landing–takeoff asymmetry is an index of a lack of elasticity, which increases with increasing the role of muscle relative to that of tendon within muscle–tendon units.     

Common motor mechanisms support body load in serially homologous legs of cockroaches in posture and walking

We studied the mechanisms underlying support of body load in posture and walking in serially homologous legs of cockroaches. Activities of the trochanteral extensor muscle in the front or middle legs were recorded neurographically while animals were videotaped. Body load was increased via magnets attached to the thorax and varied through a coil below the substrate. In posture, tonic firing of the slow trochanteral extensor motoneuron (Ds) in each leg was strongly modulated by changing body load. Rapid load increases produced decreases in body height and sharp increments in extensor firing. The peak of extensor activity more closely approximated the maximum velocity of body displacement than the body position. In walking, extensor bursts in front and middle legs were initiated during swing and continued into the stance phase. Moderate tonic increases in body load elicited similar, specific, phase dependent changes in both legs: extensor firing was not altered in swing but was higher after foot placement in stance. These motor adjustments to load are not anticipatory but apparently depend upon sensory feedback. These data are consistent with previous findings in the hind legs and support the idea that body load is countered by common motor mechanisms in serially homologous legs.     

Two Independent Contributions to Step Variability during Over-Ground Human Walking

Human walking exhibits small variations in both step length and step width, some of which may be related to active balance control. Lateral balance is thought to require integrative sensorimotor control through adjustment of step width rather than length, contributing to greater variability in step width. Here we propose that step length variations are largely explained by the typical human preference for step length to increase with walking speed, which itself normally exhibits some slow and spontaneous fluctuation. In contrast, step width variations should have little relation to speed if they are produced more for lateral balance. As a test, we examined hundreds of overground walking steps by healthy young adults (N = 14, age < 40 yrs.). We found that slow fluctuations in self-selected walking speed (2.3% coefficient of variation) could explain most of the variance in step length (59%, P < 0.01). The residual variability not explained by speed was small (1.5% coefficient of variation), suggesting that step length is actually quite precise if not for the slow speed fluctuations. Step width varied over faster time scales and was independent of speed fluctuations, with variance 4.3 times greater than that for step length (P < 0.01) after accounting for the speed effect. That difference was further magnified by walking with eyes closed, which appears detrimental to control of lateral balance. Humans appear to modulate fore-aft foot placement in precise accordance with slow fluctuations in walking speed, whereas the variability of lateral foot placement appears more closely related to balance. Step variability is separable in both direction and time scale into balance- and speed-related components. The separation of factors not related to balance may reveal which aspects of walking are most critical for the nervous system to control.     

Incoming nucleotide binds to Klenow ternary complex leading to stable physical sequestration of preceding dNTP on DNA

Klenow–DNA complex is known to undergo a rate-limiting, protein conformational transition from an ‘open’ to ‘closed’ state, upon binding of the ‘correct’ dNTP at the active site. In the ‘closed’ state, Mg²⁺ mediates a rapid chemical step involving nucleophilic displacement of pyrophosphate by the 3′ hydroxyl of the primer terminus. The enzyme returns to the ‘open’ state upon the release of PPi and translocation permits the next round of reaction. To determine whether Klenow can translocate to the next site on the addition of the next dNTP, without the preceding chemical step, we studied the ternary complex (Klenow–DNA–dNTP) in the absence of Mg²⁺. While the ternary complex is proficient in chemical addition of dNTPs in Mg²⁺, as revealed by primer extensions, the same in Mg²⁺-deficient conditions lead to non-covalent (physical) sequestration of first two ‘correct’ dNTPs in the ternary complex. Moreover, the second dNTP traps the first one in the DNA-helix of the ternary complex. Such a dNTP–DNA complex is found to be stable even after the dissociation of Klenow. This reveals the novel state of the dNTP–DNA complex where the complementary base is stacked in a DNA-helix non-covalently, without the phosphodiester linkage. Further, shuttling of the DNA between the polymerase and the exonuclease site mediates the release of such a DNA complex. Interestingly, Klenow in such a Mg²⁺-deficient ternary complex exhibits a ‘closed’ conformation.     

Estimation of stride length in level walking using an inertial measurement unit attached to the foot: a validation of the zero velocity assumption during stance

In a variety of applications, inertial sensors are used to estimate spatial parameters by double integrating over time their coordinate acceleration components. In human movement applications, the drift inherent to the accelerometer signals is often reduced by exploiting the cyclical nature of gait and under the hypothesis that the velocity of the sensor is zero at some point in stance. In this study, the validity of the latter hypothesis was investigated by determining the minimum velocity of progression of selected points of the foot and shank during the stance phase of the gait cycle while walking at three different speeds on level ground. The errors affecting the accuracy of the stride length estimation resulting from assuming a zero velocity at the beginning of the integration interval were evaluated on twenty healthy subjects. Results showed that the minimum velocity of the selected points on the foot and shank increased as gait speed increased. Whereas the average minimum velocity of the foot locations was lower than 0.011 m/s, the velocity of the shank locations were up to 0.049 m/s corresponding to a percent error of the stride length equal to 3.3%. The preferable foot locations for an inertial sensor resulted to be the calcaneus and the lateral aspect of the rearfoot. In estimating the stride length, the hypothesis that the velocity of the sensor can be set to zero sometimes during stance is acceptable only if the sensor is attached to the foot.     

Compliant bipedal model with the center of pressure excursion associated with oscillatory behavior of the center of mass reproduces the human gait dynamics

Although the compliant bipedal model could reproduce qualitative ground reaction force (GRF) of human walking, the model with a fixed pivot showed overestimations in stance leg rotation and the ratio of horizontal to vertical GRF. The human walking data showed a continuous forward progression of the center of pressure (CoP) during the stance phase and the suspension of the CoP near the forefoot before the onset of step transition. To better describe human gait dynamics with a minimal expense of model complexity, we proposed a compliant bipedal model with the accelerated pivot which associated the CoP excursion with the oscillatory behavior of the center of mass (CoM) with the existing simulation parameter and leg stiffness. Owing to the pivot acceleration defined to emulate human CoP profile, the arrival of the CoP at the limit of the stance foot over the single stance duration initiated the step-to-step transition. The proposed model showed an improved match of walking data. As the forward motion of CoM during single stance was partly accounted by forward pivot translation, the previously overestimated rotation of the stance leg was reduced and the corresponding horizontal GRF became closer to human data. The walking solutions of the model ranged over higher speed ranges (~1.7 m/s) than those of the fixed pivoted compliant bipedal model (~1.5 m/s) and exhibited other gait parameters, such as touchdown angle, step length and step frequency, comparable to the experimental observations. The good matches between the model and experimental GRF data imply that the continuous pivot acceleration associated with CoM oscillatory behavior could serve as a useful framework of bipedal model.     

Gazing into Thin Air: The Dual-Task Costs of Movement Planning and Execution during Adaptive Gait

We examined the effect of increased cognitive load on visual search behavior and measures of gait performance during locomotion. Also, we investigated how personality traits, specifically the propensity to consciously control or monitor movements (trait movement ‘reinvestment’), impacted the ability to maintain effective gaze under conditions of cognitive load. Healthy young adults traversed a novel adaptive walking path while performing a secondary serial subtraction task. Performance was assessed using correct responses to the cognitive task, gaze behavior, stepping accuracy, and time to complete the walking task. When walking while simultaneously carrying out the secondary serial subtraction task, participants visually fixated on task-irrelevant areas ‘outside’ the walking path more often and for longer durations of time, and fixated on task-relevant areas ‘inside’ the walkway for shorter durations. These changes were most pronounced in high-trait-reinvesters. We speculate that reinvestment-related processes placed an additional cognitive demand upon working memory. These increased task-irrelevant ‘outside’ fixations were accompanied by slower completion rates on the walking task and greater gross stepping errors. Findings suggest that attention is important for the maintenance of effective gaze behaviors, supporting previous claims that the maladaptive changes in visual search observed in high-risk older adults may be a consequence of inefficiencies in attentional processing. Identifying the underlying attentional processes that disrupt effective gaze behaviour during locomotion is an essential step in the development of rehabilitation, with this information allowing for the emergence of interventions that reduce the risk of falling.     

Energetics and passive dynamics of the ankle in downhill walking

This study investigated the energetics of the human ankle during the stance phase of downhill walking with the goal of modeling ankle behavior with a passive spring and damper mechanism. Kinematic and kinetic data were collected on eight male participants while walking down a ramp with inclination varying from 0° to 8°. The ankle joint moment in the sagittal plane was calculated using inverse dynamics. Mechanical energy injected or dissipated at the ankle joint was computed by integrating the power across the duration of the stance phase. The net mechanical energy of the ankle was approximately zero for level walking and monotonically decreased (i.e., became increasingly negative) during downhill walking as the slope decreased. The indication is that the behavior of the ankle is energetically passive during downhill walking, playing a key role in dissipating energy from one step to the next. A passive mechanical model consisting of a pin joint coupled with a revolute spring and damper was fit to the ankle torque and its parameters were estimated for each downhill slope using linear regression. The passive model demonstrated good agreement with actual ankle dynamics as indicated by low root-mean-square error values. These results indicate the stance phase behavior of the human ankle during downhill walking may be effectively duplicated by a passive mechanism with appropriately selected spring and damping characteristics.     

Dynamic balance during walking adaptability tasks in individuals post-stroke

Maintaining dynamic balance during community ambulation is a major challenge post-stroke. Community ambulation requires performance of steady-state level walking as well as tasks that require walking adaptability. Prior studies on balance control post-stroke have mainly focused on steady-state walking, but walking adaptability tasks have received little attention. The purpose of this study was to quantify and compare dynamic balance requirements during common walking adaptability tasks post-stroke and in healthy adults and identify differences in underlying mechanisms used for maintaining dynamic balance. Kinematic data were collected from fifteen individuals with post-stroke hemiparesis during steady-state forward and backward walking, obstacle negotiation, and step-up tasks. In addition, data from ten healthy adults provided the basis for comparison. Dynamic balance was quantified using the peak-to-peak range of whole-body angular-momentum in each anatomical plane during the paretic, nonparetic and healthy control single-leg-stance phase of the gait cycle. To understand differences in some of the key underlying mechanisms for maintaining dynamic balance, foot placement and plantarflexor muscle activation were examined. Individuals post-stroke had significant dynamic balance deficits in the frontal plane across most tasks, particularly during the paretic single-leg-stance. Frontal plane balance deficits were associated with wider paretic foot placement, elevated body center-of-mass, and lower soleus activity. Further, the obstacle negotiation task imposed a higher balance requirement, particularly during the trailing leg single-stance. Thus, improving paretic foot placement and ankle plantarflexor activity, particularly during obstacle negotiation, may be important rehabilitation targets to enhance dynamic balance during post-stroke community ambulation.     

A shoe sole-based apparatus and method for randomly perturbing the stance phase of gait: test-retest reliability in young adults

Walking on an irregular surface is associated with an elevated risk for a fall at any age. Yet, relatively little is known about how a human responds to an unexpected underfoot perturbation during gait. This is partly due to the difficulty of generating an intermittent but repeatable, unexpected, underfoot perturbation whose size and location are precisely known. So we developed a shoe sole-embedded apparatus for randomly perturbing the stance phase of gait. Medial and lateral flaps were concealed in the soles of pairs of sandals, along with their actuators. Either flap could be deployed within 400ms in the parasagittal plane under a swing foot; this altered the resulting sagittal and frontal plane orientations of the foot during the next stance phase, whereafter the flap was retracted following toe-off for the rest of that gait trial. We tested six healthy young subjects by randomly presenting a single medial or lateral perturbation in 12 of 30 gait trials. Traditional step kinematic measures were used to evaluate the test-retest reliability of the response to the stimulus at two different walking speeds in 60 randomized trials conducted 1 week apart. The method was effective in systematically inducing an alteration in gait, reproducible across visits, as evidenced by acceptable intraclass correlation coefficients for step width, time and length. We conclude that the apparatus and method has potential for measuring the ability of humans to reject one or more unexpected underfoot perturbations during gait.     

Mechanical and metabolic determinants of the preferred step width in human walking

We studied the selection of preferred step width in human walking by measuring mechanical and metabolic costs as a function of experimentally manipulated step width (0.00-0.45L, as a fraction of leg length L). We estimated mechanical costs from individual limb external mechanical work and metabolic costs using open circuit respirometry. The mechanical and metabolic costs both increased substantially (54 and 45%, respectively) for widths greater than the preferred value (0.15-0.45L) and with step width squared (R(2) = 0.91 and 0.83, respectively). As predicted by a three-dimensional model of walking mechanics, the increases in these costs appear to be a result of the mechanical work required for redirecting the centre of mass velocity during the transition between single stance phases (step-to-step transition costs). The metabolic cost for steps narrower than preferred (0.10-0.00L) increased by 8%, which was probably as a result of the added cost of moving the swing leg laterally in order to avoid the stance leg (lateral limb swing cost). Trade-offs between the step-to-step transition and lateral limb swing costs resulted in a minimum metabolic cost at a step width of 0.12L, which is not significantly different from foot width (0.11L) or the preferred step width (0.13L). Humans appear to prefer a step width that minimizes metabolic cost.     

The influence of locomotor training on dynamic balance during steady-state walking post-stroke

Slow walking speed and lack of balance control are common impairments post-stroke. While locomotor training often improves walking speed, its influence on dynamic balance is unclear. The goal of this study was to assess the influence of a locomotor training program on dynamic balance in individuals post-stroke during steady-state walking and determine if improvements in walking speed are associated with improved balance control. Kinematic and kinetic data were collected pre- and post-training from seventeen participants who completed a 12-week locomotor training program. Dynamic balance was quantified biomechanically (peak-to-peak range of frontal plane whole-body angular-momentum) and clinically (Berg-Balance-Scale and Dynamic-Gait-Index). To understand the underlying biomechanical mechanisms associated with changes in angular-momentum, foot placement and ground-reaction-forces were quantified. As a group, biomechanical assessments of dynamic balance did not reveal any improvements after locomotor training. However, improved dynamic balance post-training, observed in a sub-group of 10 participants (i.e., Responders), was associated with a narrowed paretic foot placement and higher paretic leg vertical ground-reaction-force impulse during late stance. Dynamic balance was not improved post-training in the remaining seven participants (i.e., Non-responders), who did not alter their foot placement and had an increased reliance on their nonparetic leg during weight-bearing. As a group, increased walking speed was not correlated with improved dynamic balance. However, a higher pre-training walking speed was associated with higher gains in dynamic balance post-training. These findings highlight the importance of the paretic leg weight bearing and mediolateral foot placement in improving frontal plane dynamic balance post-stroke.     

Biped gait stabilization via foot placement

It is shown that stable biped gaits can be achieved by discrete foot placement based on feedback of information available at the time of foot placement. The model, developed by Townsend (1981, J. Biomechanics 14, p. 727) to evaluate the coordinations of torso motions, subsumes most of the salient body members and motions. The modeling yielded a generalized inverted pendulum with a movable support point which physically defines lateral foot placement. The principal result is that stable gaits can be defined by foot placements which are a linear function of the system center of mass position and velocity at the time of foot placement (only). Gaits may be 'smooth' or may have impulsive corrections to adjust the character of the motions and foot placement. Several general algorithms and specific simulations are presented, and calculations for non-impulsive gaits and impulsive corrections are presented. The model predictions are compared with published data. The predictions are sufficiently close to the data such that the general algorithms appear to be validated. Of particular interest are the non-sinusoidal character of the motions and the relatively simple algorithms. Indeed, the simplicity of the algorithms suggests the practical possibility of legged mobile robots. Accordingly, further investigation seems warranted for determining the parametric variation and control of gait. Some attention is also given to continuous-feedback control such as would exist during double-leg support and in specialized tasks such as rope walking or skating. Subsequent investigation will consider superposition of single and double leg support, although clearly the discrete gaits pose the more restrictive stability problem.     

Tool for Rapid & Easy Identification of High Risk Diabetic Foot: Validation & Clinical Pilot of the Simplified 60 Second Diabetic Foot Screening Tool

Background

Most diabetic foot amputations are caused by ulcers on the skin of the foot i.e. diabetic foot ulcers. Early identification of patients at high risk for diabetic foot ulcers is crucial. The ‘Simplified 60-Second Diabetic Foot Screening Tool’ has been designed to rapidly detect high risk diabetic feet, allowing for timely identification and referral of patients needing treatment. This study aimed to determine the clinical performance and inter-rater reliability of ‘Simplified 60 Second Diabetic Foot Screening Tool’ in order to evaluate its applicability for routine screening.

Methods and Findings

The tool was independently tested by n=12 assessors with n=18 Guyanese patients with diabetes. Inter-rater reliability was assessed by calculating Cronbach’s alpha for each of the assessment items. A minimum value of 0.60 was considered acceptable. Reliability scores of the screening tool assessment items were: ‘monofilament test’ 0.98; ‘active ulcer’ 0.97; ‘previous amputation’ 0.97; ‘previous ulcer’ 0.97; ‘fixed ankle’ 0.91; ‘deformity’ 0.87; ‘callus’ 0.87; ‘absent pulses’ 0.87; ‘fixed toe’ 0.80; ‘blisters’ 0.77; ‘ingrown nail’ 0.72; and ‘fissures’ 0.55. The item ‘stiffness in the toe or ankle’ was removed as it was observed in only 1.3% of patients. The item ‘fissures’ was also removed due to low inter-rater reliability. Clinical performance was assessed via a pilot study utilizing the screening tool on n=1,266 patients in an acute care setting in Georgetown, Guyana. In total, 48% of patients either had existing diabetic foot ulcers or were found to be at high risk for developing ulcers.

Conclusions

Clinicians in low and middle income countries such as Guyana can use the Simplified 60-Second Diabetic Screening Tool to facilitate early detection and appropriate treatment of diabetic foot ulcers. Implementation of this screening tool has the potential to decrease diabetes related disability and mortality.     

Neck muscle vibration disrupts steering of locomotion.

Neck muscle vibration was applied to human subjects to assess the influences of neck abnormal proprioceptive input on the organization and execution of gait. Subjects walked blindfolded to a previously seen target, located straight ahead at ~4 m. Vibration was applied on the right side of the neck, both during and before walking. The variables measured were length, duration, and velocity of trajectory; relative and absolute frontal errors at target; and width of walking support base. Vibration applied during locomotion produced an undershoot of target and deviation of gait trajectory toward the side opposite to vibration. Vibration applied before locomotion produced no effect on length of trajectory but slowing of velocity and nonsystematic deviation. When vibration frequency was increased, the amplitude of the nonsystematic deviation increased. Vibration applied during or before stance trials had minor effects on body sway. Vibration before stance had no effect on the position of mean center of foot pressure, whereas vibration during stance displaced it to the side opposite to the vibrated muscle. We suggest that vibration during locomotion reduces length and velocity of trajectory because of a direct action on the locomotor centers and produces trajectory deviation related to its effect on stance. Vibration before locomotion causes a major, nonsystematic deviation from the planned trajectory, possibly connected to a disorientation of the internal references.     

The Relationships between Foot Arch Volumes and Dynamic Plantar Pressure during Midstance of Walking in Preschool Children

Objectives

The purpose of this study was to examine the correlation between the foot arch volume measured from static positions and the plantar pressure distribution during walking.

Methods

A total of 27 children, two to six years of age, were included in this study. Measurements of static foot posture were obtained, including navicular height and foot arch volume in sitting and standing positions. Plantar pressure, force and contact areas under ten different regions of the foot were obtained during walking.

Results

The foot arch index was correlated (r = 0.32) with the pressure difference under the midfoot during the foot flat phase. The navicular heights and foot arch volumes in sitting and standing positions were correlated with the mean forces and pressures under the first (r = −0.296∼−0.355) and second metatarsals (r = −0.335∼−0.504) and midfoot (r = −0.331∼−0.496) during the stance phase of walking. The contact areas under the foot were correlated with the foot arch parameters, except for the area under the midfoot.

Conclusions

The foot arch index measured in a static position could be a functional index to predict the dynamic foot functions when walking. The foot arch is a factor which will influence the pressure distribution under the foot. Children with a lower foot arch demonstrated higher mean pressure and force under the medial forefoot and midfoot, and lower contact areas under the foot, except for the midfoot region. Therefore, children with flatfoot may shift their body weight to a more medial foot position when walking, and could be at a higher risk of soft tissue injury in this area.