Afferent feedback in the control of human gait.

Sensory activity contributes to motor control in two fundamentally different ways. It may mediate 'error signals' following sudden external perturbations and it may contribute to the pre-programmed motoneuronal drive. Here we review data, which illustrate these two functions of sensory feedback in relation to human walking. When ankle plantarflexors are unloaded in the stance phase there is a sudden decrease in the sensory activity in muscle and tendon afferents from the active muscles. This decrease in sensory activity results in a drop in EMG activity recorded from the soleus muscle, which demonstrates that the sensory activity contributes importantly to the activation of the muscles. Data suggests that a spinal pathway from gr. II muscle afferents is responsible for this positive feedback contribution to the motoneuronal drive during walking.When cutaneous nerves from the foot are stimulated in the early swing phase of walking a late reflex response may be observed in the tibialis anterior muscle. This reflex may help to ensure that the foot is lifted effectively over an obstacle. Data suggest that this reflex response is at least partly mediated by a transcortical reflex pathway. It seems to be important that reactions to external perturbations are integrated at a supraspinal level during human walking.     

Effect of footwear on intramuscular EMG activity of plantar flexor muscles in walking

One of the purposes of footwear is to assist locomotion, but some footwear types seem to restrict natural foot motion, which may affect the contribution of ankle plantar flexor muscles to propulsion. This study examined the effects of different footwear conditions on the activity of ankle plantar flexors during walking. Ten healthy habitually shod individuals walked overground in shoes, barefoot and in flip-flops while fine-wire electromyography (EMG) activity was recorded from flexor hallucis longus (FHL), soleus (SOL), and medial and lateral gastrocnemius (MG and LG) muscles. EMG signals were peak-normalised and analysed in the stance phase using Statistical Parametric Mapping (SPM). We found highly individual EMG patterns. Although walking with shoes required higher muscle activity for propulsion than walking barefoot or with flip-flops in most participants, this did not result in statistically significant differences in EMG amplitude between footwear conditions in any muscle (p > 0.05). Time to peak activity showed the lowest coefficient of variation in shod walking (3.5, 7.0, 8.0 and 3.4 for FHL, SOL, MG and LG, respectively). Future studies should clarify the sources and consequences of individual EMG responses to different footwear.     

Experimentally reduced hip abductor function during walking: Implications for knee joint loads

Hip and knee functions are intimately connected and reduced hip abductor function might play a role in development of knee osteoarthritis (OA) by increasing the external knee adduction moment during walking. The purpose of this study was to test the hypothesis that reduced function of the gluteus medius (GM) muscle would lead to increased external knee adduction moment during level walking in healthy subjects. Reduced GM muscle function was induced experimentally, by means of intramuscular injections of hypertonic saline that produced an intense short-term muscle pain and reduced muscle function. Isotonic saline injections were used as non-painful control. Fifteen healthy subjects performed walking trials at their self-selected walking speed before and immediately after injections, and again after 20 min of rest, to ensure pain recovery. Standard gait analyses were used to calculate three-dimensional trunk and lower extremity joint kinematics and kinetics. Surface electromyography (EMG) of the glutei, quadriceps, and hamstring muscles were also measured. The peak GM EMG activity had temporal concurrence with peaks in frontal plane moments at both hip and knee joints. The EMG activity in the GM muscle was significantly reduced by pain (?39.6%). All other muscles were unaffected. Peaks in the frontal plane hip and knee joint moments were significantly reduced during pain (?6.4% and ?4.2%, respectively). Lateral trunk lean angles and midstance hip joint adduction and knee joint extension angles were reduced by ?1°. Thus, the gait changes were primarily caused by reduced GM function. Walking with impaired GM muscle function due to pain significantly reduced the external knee adduction moment. This study challenge the notion that reduced GM function due to pain would lead to increased loads at the knee joint during level walking.     

The effects of sloped surfaces on locomotion: backward walking as a perturbation

The purpose of this study was to examine lower extremity kinetics and muscle activity during backward slope walking to clarify the relationship between joint moments and powers and muscle activity patterns observed in forward slope walking. Nine healthy volunteers walked backward on an instrumented ramp at three grades (-39% (-21 degrees ), 0% (level), +39% (+21 degrees )). EMG activity was recorded from major lower extremity muscles. Joint kinetics were obtained from kinematic and force platform data. The knee joint moment and power generation increased significantly during upslope walking; hip joint moment and power absorption increased significantly during downslope walking. When compared to data from forward slope walking, these backward walking data suggest that power requirements of a task dictate the muscle activity pattern needed to accomplish that movement. During downslope walking tasks, power absorption increased and changes in muscle activity patterns were directly related to the changes in the joint moment patterns. In contrast, during upslope walking tasks, power generation increased and changes in the muscle activity were related to the changes in the joint moments only at the 'primary' joint; at adjacent joints the changes in muscle activity were unrelated to the joint moment pattern. The 'paradoxical' changes in the muscle activity at the adjacent joints are possibly related to the activation of biarticular muscles required by the increased power generation at the primary joint. In total, these data suggest that changing power requirements at a joint impact the control of muscle activity at that and adjacent joints.     

The influence of footwear on the electromyographic activity of selected lower limb muscles during walking

The purpose of this study was to compare the effects of a standard flexible shoe and a stability running shoe on lower limb muscle activity during walking. Twenty-eight young asymptomatic adults with flat-arched feet were recruited. While walking, electromyographic (EMG) activity was recorded from tibialis posterior and peroneus longus via intramuscular electrodes; and from tibialis anterior and medial gastrocnemius via surface electrodes. Three experimental conditions were assessed: (i) barefoot, (ii) a standard flexible shoe, (iii) a stability running shoe. Results showed significant differences for the peak amplitude and the time of peak amplitude for tibialis anterior, peroneus longus and medial gastrocnemius when comparing the three experimental conditions (p < 0.05). Significant differences were detected primarily between the barefoot and shoe conditions and with relatively small effect sizes for peroneus longus, tibialis anterior and medial gastrocnemius. Few significant differences were found between the two shoe styles. We discuss how these changes are most likely associated with the shoe upper bracing the foot, the shape of the shoe outer-sole and weight of the shoes. Further research is needed to investigate differences between these shoe styles when participants walk for longer distances (i.e. over 1000 m) and following fatigue.     

A study of the viability of obtaining a generic animation of the foot while walking for the virtual testing of footwear using dorsal pressures

Establishing the appropriate pressure exerted by the shoe upper over the foot surface is fundamental for the design of specific footwear, although measuring the dorsal pressures can also provide important additional information. In previous works, a virtual simulator to perform studies of comfort and functionality in CAD footwear design was presented. This paper describes the procedure carried out to obtain the foot animations used in this simulator. The virtual feet used in the simulator are feet without a standard form scanned in a static way. Their movements are rebuilt from the register of movements of several foot anatomical points during a complete step. The dorsal pressures exerted by some shoe uppers on these anatomical points were measured for several subjects and used to establish the viability of the use of these animations in a virtual simulator for footwear.     

Trunk kinematic,kinetic, and neuro-muscular response to foot center of pressure translation along the medio-lateral foot axis during gait

Footwear devices that shift foot center of pressure (COP), thereby impacting lower-limb biomechanics to produce clinical benefit, have been studied regarding degenerative diseases of knee and hip joints, exhibiting evidence of clinical success. Ability to purposefully affect trunk biomechanics has not been investigated for this type of footwear. Fifteen healthy young male subjects underwent gait and electromyography analysis using a biomechanical device that shifts COP via moveable convex elements attached to the shoe sole. Analyses were performed in three COP configurations for pairwise comparison: (1) neutral (control) (2) laterally deviated, and (3) medially deviated. Sagittal and frontal-plane pelvis and spine kinematics, external oblique activity, and frontal and transverse-plane lumbar moments were affected by medio-lateral COP shift. Transverse-plane trunk kinematics, activity of the lumbar longissimus, latissimus dorsi, rectus abdominus, and quadratus lumborum, and sagittal-plane lumbar moment, were not significantly impacted. Two linear mixed effects models assessed predictive impact of (I) COP location, and (II) trunk kinematics and neuromuscular activity, on the significant lumbar moment parameters. The COP was a significant predictor of all modeled frontal and transverse-plane lumbar moment parameters, while pelvic and spine rotation, and lumbar longissimus activity were significant predictors of one frontal-plane lumbar moment parameter. Model results suggest that, although trunk biomechanics and muscle activity were altered by COP shift, COP offset influences lumbar kinetics directly, or via lower-limb changes not assessed in this study, but not by means of alteration of trunk kinematics or muscle activity. Further study may reveal implications in treatment of low back pain.     

Skeletal kinematics of the midtarsal joint during walking: Midtarsal joint locking revisited

The kinematics of the human foot complex have been investigated to understand the weight bearing mechanism of the foot. This study aims to investigate midtarsal joint locking during walking by noninvasively measuring the movements of foot bones using a high-speed bi-planar fluoroscopic system. Eighteen healthy subjects volunteered for the study; the subjects underwent computed tomography imaging and bi-planar radiographs of the foot in order to measure the three-dimensional (3D) midtarsal joint kinematics using a 2D-to-3D registration method and anatomical coordinate system in each bone. The relative movements on bone surfaces were also calculated in the talonavicular and calcaneocuboid joints and quantified as surface relative velocity vectors on articular surfaces to understand the kinematic interactions in the midtarsal joint. The midtarsal joint performed a coupled motion in the early stance to pronate the foot to extreme pose in the range of motion during walking and maintained this pose during the mid-stance. In the terminal stance, the talonavicular joint performed plantar-flexion, inversion, and internal rotation while the calcaneocuboid joint performed mainly inversion. The midtarsal joint moved towards an extreme supinated pose, rather than a minimum motion in the terminal stance. The study provides a new perspective to understand the kinematics and kinetics of the movement of foot bones and so-called midtarsal joint locking, during walking. The midtarsal joint continuously moved towards extreme poses together with the activation of muscle forces, which would support the foot for more effective force transfer during push-off in the terminal stance.     

In vitro study of foot kinematics using a dynamic walking cadaver model

There is a dearth of information on navicular, cuboid, cuneiform and metatarsal kinematics during walking and our objective was to study the kinematic contributions these bones might make to foot function. A dynamic cadaver model of walking was used to apply forces to cadaver feet and mobilise them in a manner similar to in vivo. Kinematic data were recorded from 13 cadaver feet. Given limitations to the simulation, the data describe what the cadaver feet were capable of in response to the forces applied, rather than exactly how they performed in vivo. The talonavicular joint was more mobile than the calcaneocuboid joint. The range of motion between cuneiforms and navicular was similar to that between talus and navicular. Metatarsals four and five were more mobile relative to the cuboid than metatarsals one, two and three relative to the cuneiforms. This work has confirmed the complexity of rear, mid and forefoot kinematics. The data demonstrate the potential for often-ignored foot joints to contribute significantly to the overall kinematic function of the foot. Previous emphasis on the ankle and sub talar joints as the principal articulating components of the foot has neglected more distal articulations. The results also demonstrate the extent to which the rigid segment assumptions of previous foot kinematics research have over simplified the foot.     

Unholey shoes: Experimental considerations when estimating ankle joint complex power during walking and running

For studies that aim to assess biological ankle function, calculating ankle joint complex (AJC) power between the calcaneus and shank is recommended over conventional inverse dynamics estimates between a rigid-body foot and shank. However, when designing a new experiment, it remains unclear whether holes should be cut in footwear to permit motion tracking via skin-mounted markers, or whether marker placement locations should be tightly controlled across conditions. Here we provide data to assist researchers in answering these questions. We performed a gait analysis study of walking (0.8, 1.2, 1.6 m·s⁻¹) and running (2.6, 2.8, 3.0 m·s⁻¹) while subjects (N = 10) wore custom-modified footwear, which allowed markers to be placed either on the shoe, or on the skin via cut-out windows in the shoes. First, we compared foot markers affixed to the skin vs. on the same locations on the shoe. Using statistical non-parametric mapping techniques, we discovered that skin vs. shoe markers had no statistically significant effect on net AJC power estimates throughout stance phase, for all walking and running speeds. Second, we compared calcaneal markers in the nominal shoe configuration vs. markers in a nearby location (∼27 mm below) on the shoe. We observed significant differences when marker placement on the shoe was varied, which may be relevant to repeated-measures study designs. The results suggest that when computing AJC power for walking and running, you may want to put down the scissors (i.e., forego cutting holes in your footwear), and instead pick up a Sharpie® (pen) or use a template, to maintain consistent marker placement across trials and conditions.     

Walking with increased ankle pushoff decreases hip muscle moments

In a simple bipedal walking model, an impulsive push along the trailing limb (similar to ankle plantar flexion) or a torque at the hip can power level walking. This suggests a tradeoff between ankle and hip muscle requirements during human gait. People with anterior hip pain may benefit from walking with increased ankle pushoff if it reduces hip muscle forces. The purpose of our study was to determine if simple instructions to alter ankle pushoff can modify gait dynamics and if resulting changes in ankle pushoff have an effect on hip muscle requirements during gait. We hypothesized that changes in ankle kinetics would be inversely related to hip muscle kinetics. Ten healthy subjects walked on a custom split-belt force-measuring treadmill at 1.25m/s. We recorded ground reaction forces and lower extremity kinematic data to calculate joint angles and internal muscle moments, powers and angular impulses. Subjects walked under three conditions: natural pushoff, decreased pushoff and increased pushoff. For the decreased pushoff condition, subjects were instructed to push less with their feet as they walked. Conversely, for the increased pushoff condition, subjects were instructed to push more with their feet. As predicted, walking with increased ankle pushoff resulted in lower peak hip flexion moment, power and angular impulse as well as lower peak hip extension moment and angular impulse (p<0.05). Our results emphasize the interchange between hip and ankle kinetics in human walking and suggest that increased ankle pushoff during gait may help to compensate for hip muscle weakness or injury and reduce hip joint forces.     

Examination of extrinsic foot muscles during running using mfMRI and EMG.

Over-pronation has been cited as a key contributor to many types of running injuries. However, the roles of the extrinsic foot muscles during running have not been adequately identified. The purpose of this study was to examine the muscle functional (mf) MRI and EMG responses to perturbations of the foot by running in varus, neutral and valgus wedged shoes. Ten males ran at 3.6 m/s in specially constructed shoes for 5 min with T2-weighted mfMRI collected before and after each run. The change in T2 from before to after each run characterized the level of metabolic activity in each of muscle. Kinematic and EMG data were also collected while subjects ran on a treadmill. There were no T2 differences across the three shoe conditions. In contrast, there was significantly less EMG activity in the tibialis anterior and soleus while wearing the neutral shoe. Overall, the results did not support the theory that muscle activity would increase as the degree of eversion increased. It also appears that surface EMG was more sensitive to differences between conditions than mfMRI. However, this study illustrated that mfMRI may be a useful tool for quantifying muscle activity in cases where surface EMG is inadequate.     

The free moment of ground reaction in distance running and its changes with pronation.

Many running injuries are successfully treated with footwear modifications designed to reduce pronation, but the underlying mechanism of treatment is not well understood. Previous attempts to correlate reduction in pronation with changes in ground reaction parameters have been unsuccessful. In this study, the free moment of ground reaction (Mz') was measured for 10 rearfoot strikers running at 4.5 m s-1 in each of three different pairs of running shoes designed to vary the extent of pronation during ground contact. Mz' patterns were highly variable between feet, but were repeatable within a given foot/footwear combination. Mz' was greatest in magnitude during the first half of support, when it acted in a direction resisting foot abduction, a component of pronation. It was opposite in sign and smaller in magnitude during the last 30% of support. The peak magnitude and the net angular impulse of Mz' were both increased significantly with increases in pronation. A net ground reaction moment was also calculated about a vertical axis fixed in the shoe, and was used in a first approximation model of the shoe/ground interface to predict when the foot is most likely to ab/adduct during running. In conclusion, this study characterized the Mz' pattern for a well-defined group of runners, and found that Mz' is sensitive to relatively large within-subject changes in pronation.     

Predicting the metabolic cost of incline walking from muscle activity and walking mechanics

The goal of this study was to identify which muscle activation patterns and gait features best predict the metabolic cost of inclined walking. We measured muscle activation patterns, joint kinematics and kinetics, and metabolic cost in sixteen subjects during treadmill walking at inclines of 0%, 5%, and 10%. Multivariate regression models were developed to predict the net metabolic cost from selected groups of the measured variables. A linear regression model including incline and the squared integrated electromyographic signals of the soleus and vastus lateralis explained 96% of the variance in metabolic cost, suggesting that the activation patterns of these large muscles have a high predictive value for metabolic cost. A regression model including only the peak knee flexion angle during stance phase, peak knee extension moment, peak ankle plantarflexion moment, and peak hip flexion moment explained 89% of the variance in metabolic cost; this finding indicates that kinematics and kinetics alone can predict metabolic cost during incline walking. The ability of these models to predict metabolic cost from muscle activation patterns and gait features points the way toward future work aimed at predicting metabolic cost when gait is altered by changes in neuromuscular control or the use of an assistive technology.     

Response of able-bodied persons to changes in shoe rocker radius during walking: Changes in ankle kinematics to maintain a consistent roll-over shape

Recent studies have determined a seemingly consistent feature of able-bodied level ground walking, termed the roll-over shape, which is the effective rocker (cam) shape that the lower limb system conforms to between heel contact and contralateral heel contact during walking (first half of the gait cycle). The roll-over shape has been found to be largely unaffected by changes in walking speed, load carriage, and shoe heel height. However, it is unclear from previous studies whether persons are controlling their lower limb systems to maintain a consistent roll-over shape or whether this finding is a byproduct of their attempt to keep ankle kinematic patterns similar during the first half of the gait cycle. We measured the ankle–foot roll-over shapes and ankle kinematics of eleven able-bodied subjects while walking on rocker shoes of different radii. We hypothesized that the ankle flexion patterns during single support would change to maintain a similar roll-over shape. We also hypothesized that with decrease in rocker shoe radii, the difference in ankle flexion between the end and beginning of single support would decrease. Our results supported these hypotheses. Ankle kinematics were changed significantly during walking with the different rocker shoe radii (p<0.001), while ankle–foot roll-over shape radii (p=0.146) and fore–aft position (p=0.132) were not significantly affected. The results of this study have direct implications for designers of ankle–foot prostheses, orthoses, walking casts/boots, and rocker shoes. The results may also be relevant to researchers studying control of human movements.     

Changes in muscle activity in response to different impact forces affect soft tissue compartment mechanical properties

Electromyographic (EMG) activity is associated with several tasks prior to landing in walking and running including positioning the leg, developing joint stiffness and possibly control of soft tissue compartment vibrations. The concept of muscle tuning suggests one reason for changes in muscle activity pattern in response to small changes in impact conditions, if the frequency content of the impact is close to the natural frequency of the soft tissue compartments, is to minimize the magnitude of soft tissue compartment vibrations. The mechanical properties of the soft tissue compartments depend in part on muscle activations and thus it was hypothesized that changes in the muscle activation pattern associated with different impact conditions would result in a change in the acceleration transmissibility to the soft tissue compartments. A pendulum apparatus was used to systematically administer impacts to the heel of shod male participants. Wall reaction forces, EMG of selected leg muscles, soft tissue compartment and shoe heel cup accelerations were quantified for two different impact conditions. The transmissibility of the impact acceleration to the soft tissue compartments was determined for each subject/soft tissue compartment/shoe combination. For this controlled impact situation it was shown that changes in the damping properties of the soft tissue compartments were related to changes in the EMG intensity and/or mean frequency of related muscles in response to a change in the impact interface conditions. These results provide support for the muscle tuning idea--that one reason for the changes in muscle activity in response to small changes in the impact conditions may be to minimize vibrations of the soft tissue compartments that are initiated at heel-strike.     

Comparison of Trunk Activity during Gait Initiation and Walking in Humans

To understand the role of trunk muscles in maintenance of dynamic postural equilibrium we investigate trunk movements during gait initiation and walking, performing trunk kinematics analysis, Erector spinae muscle (ES) recordings and dynamic analysis. ES muscle expressed a metachronal descending pattern of activity during walking and gait initiation. In the frontal and horizontal planes, lateroflexion and rotation occur before in the upper trunk and after in the lower trunk. Comparison of ES muscle EMGs and trunk kinematics showed that trunk muscle activity precedes corresponding kinematics activity, indicating that the ES drive trunk movement during locomotion and thereby allowing a better pelvis mobilization. EMG data showed that ES activity anticipates propulsive phases in walking with a repetitive pattern, suggesting a programmed control by a central pattern generator. Our findings also suggest that the programs for gait initiation and walking overlap with the latter beginning before the first has ended.     

Similar sensorimotor transformations control balance during standing and walking

Standing and walking balance control in humans relies on the transformation of sensory information to motor commands that drive muscles. Here, we evaluated whether sensorimotor transformations underlying walking balance control can be described by task-level center of mass kinematics feedback similar to standing balance control. We found that delayed linear feedback of center of mass position and velocity, but not delayed linear feedback from ankle angles and angular velocities, can explain reactive ankle muscle activity and joint moments in response to perturbations of walking across protocols (discrete and continuous platform translations and discrete pelvis pushes). Feedback gains were modulated during the gait cycle and decreased with walking speed. Our results thus suggest that similar task-level variables, i.e. center of mass position and velocity, are controlled across standing and walking but that feedback gains are modulated during gait to accommodate changes in body configuration during the gait cycle and in stability with walking speed. These findings have important implications for modelling the neuromechanics of human balance control and for biomimetic control of wearable robotic devices. The feedback mechanisms we identified can be used to extend the current neuromechanical models that lack balance control mechanisms for the ankle joint. When using these models in the control of wearable robotic devices, we believe that this will facilitate shared control of balance between the user and the robotic device.     

Can Treadmill Perturbations Evoke Stretch Reflexes in the Calf Muscles?

Disinhibition of reflexes is a problem amongst spastic patients, for it limits a smooth and efficient execution of motor functions during gait. Treadmill belt accelerations may potentially be used to measure reflexes during walking, i.e. by dorsal flexing the ankle and stretching the calf muscles, while decelerations show the modulation of reflexes during a reduction of sensory feedback. The aim of the current study was to examine if belt accelerations and decelerations of different intensities applied during the stance phase of treadmill walking can evoke reflexes in the gastrocnemius, soleus and tibialis anterior in healthy subjects. Muscle electromyography and joint kinematics were measured in 10 subjects. To determine whether stretch reflexes occurred, we assessed modelled musculo-tendon length and stretch velocity, the amount of muscle activity, as well as the incidence of bursts or depressions in muscle activity with their time delays, and co-contraction between agonist and antagonist muscle. Although the effect on the ankle angle was small with 2.8±1.0°, the perturbations caused clear changes in muscle length and stretch velocity relative to unperturbed walking. Stretched muscles showed an increasing incidence of bursts in muscle activity, which occurred after a reasonable electrophysiological time delay (163–191 ms). Their amplitude was related to the muscle stretch velocity and not related to co-contraction of the antagonist muscle. These effects increased with perturbation intensity. Shortened muscles showed opposite effects, with a depression in muscle activity of the calf muscles. The perturbations only slightly affected the spatio-temporal parameters, indicating that normal walking was retained. Thus, our findings showed that treadmill perturbations can evoke reflexes in the calf muscles and tibialis anterior. This comprehensive study could form the basis for clinical implementation of treadmill perturbations to functionally measure reflexes during treadmill-based clinical gait analysis.