Contribution of sensory feedback to ongoing ankle extensor activity during the stance phase of walking

Numerous investigations over the past 15 years have demonstrated that sensory feedback plays a critical role in establishing the timing and magnitude of muscle activity during walking. Here we review recent studies reporting that sensory feedback makes a substantial contribution to the activation of extensor motoneurons during the stance phase. Quantitative analysis of the effects of loading and unloading ankle extensor muscles during walking on a horizontal surface has shown that sensory feedback can increase the activity of ankle extensor muscles by up to 60%. There is currently some uncertainty about which sensory receptors are responsible for this enhancement of extensor activity, but likely candidates are the secondary spindle endings in the ankle extensors of humans and the Golgi tendon organs in the ankle extensors of humans and cats. Two important issues arise from the finding that sensory feedback from the leg regulates the magnitude of extensor activity. The first is the extent to which differences in the magnitude of activity in extensor muscles during different locomotor tasks can be directly attributed to changes in the magnitude of sensory signals, and the second is whether the enhancement of extensor activity is determined primarily by feedback from a specific group of receptors or from numerous groups of receptors distributed throughout the leg. Limitations of current experimental strategies prevent a straightforward empirical resolution of these issues. A potentially fruitful approach in the immediate future is to develop models of the known and hypothesized neuronal networks controlling motoneuronal activity, and use these simulations to control forward dynamic models of the musculo-skeletal system. These simulations would help understand how sensory signals are modified with a change in locomotor task and, in conjunction with physiological experiments, establish the extent to which these modifications can account for changes in the magnitude of motoneuronal activity.     

Talocrural and talocalcaneal joint kinematics and kinetics during the stance phase of walking

The purpose of this investigation was to study the kinematics and kinetics of the joints between the leg and calcaneus during the stance phase of walking. The talocrural and talocalcaneal joints were each assumed to act as monocentric single degree of freedom hinge joints. Motion at one joint was defined by the relative rotation of a point on the opposing joint. The results, based upon the gait of three subjects, showed that the hinge joint assumption may be reasonable. A discrepancy in the kinematics was shown between the talocrural joint rotation and its commonly assumed sagittal plane representation, especially during initial flatfoot. This discrepancy is due to the fact that the sagittal plane rotation is created by the combined rotations of the talocrural and talocalcaneal joints. The talocalcaneal joint showed a peak 25-30 Nm supinatory moment at 80% of stance. The talocrural joint moment was qualitatively similar to the commonly measured sagittal plane moment, but the present results show that the sagittal plane moment overpredicted the true moment by 6-22% due to the two-dimensional assumption.     

Selective activation of human soleus and medial gastrocnemius muscles during walking in water

During walking in water (WW) the vertical component of ground reaction forces decreases, while the greater propulsive force is required to move forward against the greater resistance of water. In such reduced gravity environment, Hutchison et al. (1989) have demonstrated that the relative activation of rat medial gastrocnemius (MGAS) increased compared to that of the soleus (SOL) during swimming, suggesting different effects of peripheral information on motoneuron excitability of these muscles. It is conceivable that both buoyancy and resistance of water have different effects on the activation patterns of triceps surae muscles during WW, since the reduced weight in water might decrease the peripheral inflow relating load information while greater volitional command might be needed to propel a body forward against the water resistance. The present study was designed to assess each peripheral inflow and efferent input by adjusting the load and walking speed voluntarily during WW. The aim of this study is to investigate the dissociative activation pattern between the SOL and the MGAS during WW.     

Distinct fast and slow processes contribute to the selection of preferred step frequency during human walking

Humans spontaneously select a step frequency that minimizes the energy expenditure of walking. This selection might be embedded within the neural circuits that generate gait so that the optimum is pre-programmed for a given walking speed. Or perhaps step frequency is directly optimized, based on sensed feedback of energy expenditure. Direct optimization is expected to be slow due to the compounded effect of delays and iteration, whereas a pre-programmed mechanism presumably allows for faster step frequency selection, albeit dependent on prior experience. To test for both pre-programmed selection and direct optimization, we applied perturbations to treadmill walking to elicit transient changes in step frequency. We found that human step frequency adjustments (n = 7) occurred with two components, the first dominating the response (66 ± 10% of total amplitude change; mean ± SD) and occurring quite quickly (1.44 ± 1.14 s to complete 95% of total change). The other component was of smaller amplitude (35 ± 10% of total change) and took tens of seconds (27.56 ± 16.18 s for 95% completion). The fast process appeared to be too fast for direct optimization and more indicative of a pre-programmed response. It also persisted even with unusual closed-loop perturbations that conflicted with prior experience and rendered the response energetically suboptimal. The slow process was more consistent with the timing expected for direct optimization. Our interpretation of these results is that humans may rely heavily on pre-programmed gaits to rapidly select their preferred step frequency and then gradually fine-tune that selection with direct optimization.     

All leg joints contribute to quiet human stance: A mechanical analysis

According to the state of the art model (single inverted pendulum) the regulation of quiet human stance seems to be dominated by ankle joint actions. Recent findings substantiated both in-phase and anti-phase fluctuations of ankle and hip joint kinematics can be identified in quiet human stance. Thus, we explored in an experimental study to what extent all three leg joints actually contribute to the balancing problem of quiet human stance. We also aimed at distinguishing kinematic from torque contributions. Thereto, we directly measured ankle, knee, and hip joint kinematics with high spatial resolution and ground reaction forces. Then, we calculated the six respective joint torques and, additionally, the centre of mass kinematics. We searched for high cross-correlations between all these mechanical variables. Beyond confirming correlated anti-phase kinematics of ankle and hip, the main results are: (i) ankle and knee joint fluctuate tightly (torque) coupled and (ii) the bi-articular muscles of the leg are well suited to fulfil the requirements of fluctuations around static equilibrium. Additionally, we (iii) identified high-frequency oscillations of the shank between about 4 and 8 Hz and (iv) discriminated potentially passive and active joint torque contributions. These results demonstrate that all leg joints contribute actively and concertedly to quiet human stance, even in the undisturbed case. Moreover, they substantiate the single inverted pendulum paradigm to be an invalid model for quiet human stance.     

Reliability of an individually molded shank shell for measuring tibial transverse rotations during the stance phase of walking

Use of a shank shell has been shown to estimate tibial transverse rotations better than skin-mounted markers. However, the day-to-day reliability of the transverse tibial rotations using an individually molded shank shell has not been previously investigated. This study examined the between-tests and trials reliability of an individually molded shank shell for measuring peak tibial internal and external rotations, time of peak values, and tibia range of motion during 5 walking trials. The trial-to-trial reliability of tibial transverse rotations was measured in 14 healthy individuals while the test-retest reliability was measured in 10 persons on two occasions. Trial-to-trial reliability for peak transverse rotations, time of peak values, and tibia range of motion ranged from ICC (3,1) 0.59-0.95. The PCA between trials showed that 88-99 % of values were within 3 degrees of agreement. Test-retest reliability for peak rotations, tibia range of motion, and time of peak values ranged from ICC (3,1) 0.70-0.89 with SEM 1.6-2.21 degrees , 0.021 %, and 0.034 %, respectively. The PCA between tests showed that 70-100 % of values were within 3 degrees of agreement. The use of an individually molded shell and the close attachment of the shank shell to the individual's shank resulted in reliable test-retest and trial-to-trial data.     

Empirical evaluation of gastrocnemius and soleus function during walking

Distinguishing gastrocnemius and soleus muscle function is relevant for treating gait disorders in which abnormal plantarflexor activity may contribute to pathological movement patterns. Our objective was to use experimental and computational analysis to determine the influence of gastrocnemius and soleus activity on lower limb movement, and determine if anatomical variability of the gastrocnemius affected its function. Our hypothesis was that these muscles exhibit distinct functions, with the gastrocnemius inducing limb flexion and the soleus inducing limb extension. To test this hypothesis, the gastrocnemius or soleus of 20 healthy participants was electrically stimulated for brief periods (90 ms) during mid- or terminal stance of a random gait cycle. Muscle function was characterized by the induced change in sagittal pelvis, hip, knee, and ankle angles occurring during the 200 ms after stimulation onset. Results were corroborated with computational forward dynamic gait models, by perturbing gastrocnemius or soleus activity during similar portions of the gait cycle. Mid- and terminal stance gastrocnemius stimulation induced posterior pelvic tilt, hip flexion and knee flexion. Mid-stance gastrocnemius stimulation also induced ankle dorsiflexion. In contrast mid-stance soleus stimulation induced anterior pelvic tilt, knee extension and plantarflexion, while late-stance soleus stimulation induced relatively little change in motion. Model predictions of induced hip, knee, and ankle motion were generally in the same direction as those of the experiments, though the gastrocnemius? results were shown to be quite sensitive to its knee-to-ankle moment arm ratio.     

The between-day reliability of peroneus longus EMG during walking

The peroneus longus (PL) is a rearfoot evertor, important in frontal plane foot motion. Studying PL function has been limited by previous electromyography (EMG) studies reporting poor between-day reliability. Due to its close proximity to adjacent muscles, EMG measures of PL may be susceptible to crosstalk, thus correct electrode placement is vital. The aim of this study was to use ultrasound to aid placement of small surface EMG electrodes and determine the between-day reliability of PL EMG in healthy participants’ walking. Ten participants walked barefoot and shod at a controlled, self-selected speed. Six trials per condition, per session, were recorded over two days (mean (SD): 5 (3) days apart). The muscle belly was located using ultrasound. EMG was recorded with surface electrodes (Trigno™ Mini, Delsys, Inc.) at 2000 Hz. Amplitude was normalized to the peak per gait cycle and time normalized to the gait cycle. Reliability of discrete variables were primarily assessed with the standard error of measurement (SEM), plus the coefficient of multiple correlation (CMC), the coefficient of variation (CV) and the intra-class correlation coefficient (ICC). The pattern of the EMG profile was consistent. The SEM of peak amplitude was 4% (3–8%) and 3% (2–5%) for barefoot and shod respectively. For timing of the peak the SEM was 2% (1–3%) and 1% (1–2%) for barefoot and shod respectively. Low SEM of discrete variables suggests good reliability of PL EMG during walking supporting the future use of this protocol. Therefore activation of PL can be confidently studied in repeated-measures study designs.     

In vivo biomechanical behavior of the human heel pad during the stance phase of gait

A technique is introduced for simultaneous measurements of the heel pad tissue deformation and the heel–ground contact stresses developing during the stance phase of gait. Subjects walked upon a gait platform integrating the contact pressure display optical method for plantar pressure measurements and a digital radiographic fluoroscopy system for skeletal and soft tissue motion recording. Clear images of the posterior-plantar aspect of the calcaneus and enveloping soft tissues were obtained simultaneously with the pressure distribution under the heel region throughout the stance phase of gait. The heel pad was shown to undergo a rapid compression during initial contact and heel strike, reaching a strain of 0.39±0.05 in about 150 ms. The stress–strain relation of the heel pad was shown to be highly non-linear, with a compression modulus of 105±11 kPa initially and 306±16 kPa at 30% strain. The energy dissipation during heel strike was evaluated to be 17.8±0.8%. The present technique is useful for biomechanical as well as clinical evaluation of the stress–strain and energy absorption characteristics of the heel pad in vivo, during natural gait.     

Using biofeedback to reduce left arm extensor EMG of string players during musical performance

Electromyographic (EMG) feedback offers a mechanism for helping musicians reduce specific muscle tension during performance. Nine intermediate to advanced level string players participated in a four-session, pretest/posttest design study to determine (1) if left forearm extensor EMG could be reduced using biofeedback, (2) if reductions in EMG would generalize to a no-feedback condition, and (3) if reductions in EMG would generalize from extensors to flexors. Results indicate that biofeedback did facilitate significant decreases in EMG, that the reductions in EMG did generalize to a no-feedback condition, and that generalization from extensors to flexors did not occur.     

Diabetic neuropathy is related to joint stiffness during late stance phase

The majority of plantar ulcers in the diabetic population occur in the forefoot. Peripheral neuropathy has been related to the occurrence of ulcers. Long-term diabetes results in the joints becoming passively stiffer. This static stiffness may translate to dynamic joint stiffness in the lower extremities during gait. Therefore, the purpose of this investigation was to demonstrate differences in ankle and knee joint stiffness between diabetic individuals with and without peripheral neuropathy during gait. Diabetic subjects with and without peripheral neuropathy were compared. Subjects were monitored during normal walking with three-dimensional motion analysis and a force plate. Neuropathic subjects had higher ankle stiffness (0.236 N.m/deg) during 65 to 80% of stance when compared with non-neuropathic subjects (-0.113 N.m/deg). Neuropathic subjects showed a different pattern in ankle stiffness compared with non-neuropathic subjects. Neuropathic subjects demonstrated a consistent level of ankle stiffness, whereas non-neuropathic subjects showed varying levels of stiffness. Neuropathic subjects demonstrated lower knee stiffness (0.015 N.m/deg) compared with non-neuropathic subjects (0.075 N.m/deg) during 50 to 65% of stance. The differences in patterns of ankle and knee joint stiffness between groups appear to be related to changes in timing of peak ankle dorsiflexion during stance, with the neuropathic group reaching peak dorsiflexion later than the non-neuropathic subjects. This may partially relate to the changes in plantar pressures beneath the metatarsal heads present in individuals with neuropathy.     

EMG activity of low back extensor muscles during cyclic flexion/extension

Cyclic trunk flexion/extension is known to be a risk factor for low back pain or disorders. Changes in the in vivo low back musculature associated with cyclic flexion/extension have not been adequately examined. The purpose of this study was to investigate the effects of cyclic flexion/extension on muscle activity of the low back extensors by quantifying changes in activation timing and mean amplitude of electromyography (EMG). Trunk flexion angle and EMG were recorded during 30 cyclic flexion/extension movements over a 5 min period, and during isometric extension performed before and after the 5 min period. Mean EMG in isometric extension increased (30% increase in average) significantly after cyclic flexion/extension, confirming a transfer of extension moment from viscoelastic passive tissues to the extensor muscles in isometric extension. However, in the extension phase of cyclic flexion/extension, a significant delay in the re-activation and a decrease in the mean amplitude of EMG were observed. The results of this study, together with findings in previous research, suggest that the biomechanical effects and potential risk associated with cyclic flexion/extension may be sensitive to the task demands. Further research is needed to investigate how different moment generating components function together to compensate for the loss of tissue stiffness under varying task conditions.     

Between-day repeatability of lower limb EMG measurement during running and walking

There are minimal data describing the between-day repeatability of EMG measurements during running. Furthermore, there are no data characterising the repeatability of surface EMG measurement from the adductor muscles, during running or walking. The purpose of this study was to report on the consistency of EMG measurement for both running and walking across a comprehensive set of lower limb muscles, including adductor magnus, longus and gracilis. Data were collected from 12 lower limb muscles during overground running and walking on two separate days. The coefficient of multiple correlation (CMC) was used to quantify waveform similarity across the two sessions for signals normalised to either maximal voluntary isometric contraction (MVIC) or mean/peak signal magnitude. For running, the data showed good or excellent repeatability (CMC = 0.87–0.96) for all muscles apart from gracilis and biceps femoris using the MVIC method. Similar levels of repeatability were observed for walking. Importantly, using the peak/mean method as an alternative to the MVIC method, resulted in only marginal improvements in repeatability. The proposed protocol facilitated the collection of repeatable EMG data during running and walking and therefore could be used in future studies investigating muscle patterns during gait.     

In vivo tibiofemoral cartilage deformation during the stance phase of gait

The knowledge of articular cartilage contact biomechanics in the knee joint is important for understanding the joint function and cartilage pathology. However, the in vivo tibiofemoral articular cartilage contact biomechanics during gait remains unknown. The objective of this study was to determine the in vivo tibiofemoral cartilage contact biomechanics during the stance phase of treadmill gait. Eight healthy knees were magnetic resonance (MR) scanned and imaged with a dual fluoroscopic system during gait on a treadmill. The tibia, femur and associated cartilage were constructed from the MR images and combined with the dual fluoroscopic images to determine in vivo cartilage contact deformation during the stance phase of gait. Throughout the stance phase of gait, the magnitude of peak compartmental contact deformation ranged between 7% and 23% of the resting cartilage thickness and occurred at regions with thicker cartilage. Its excursions in the anteroposterior direction were greater in the medial tibiofemoral compartment as compared to those in the lateral compartment. The contact areas throughout the stance phase were greater in the medial compartment than in the lateral compartment. The information on in vivo tibiofemoral cartilage contact biomechanics during gait could be used to provide physiological boundaries for in vitro testing of cartilage. Also, the data on location and magnitude of deformation among non-diseased knees during gait could identify where loading and later injury might occur in diseased knees.     

Biomechanical analysis of the stance phase during barefoot and shod running

This study investigated spatio-temporal variables, ground reaction forces and sagittal and frontal plane kinematics during the stance phase of nine trained subjects running barefoot and shod at three different velocities (3.5, 4.5, 5.5 m s(-1)). Differences between conditions were detected with the general linear method (factorial model). Barefoot running is characterized by a significantly larger external loading rate than the shod condition. The flatter foot placement at touchdown is prepared in free flight, implying an actively induced adaptation strategy. In the barefoot condition, plantar pressure measurements reveal a flatter foot placement to correlate with lower peak heel pressures. Therefore, it is assumed that runners adopt this different touchdown geometry in barefoot running in an attempt to limit the local pressure underneath the heel. A significantly higher leg stiffness during the stance phase was found for the barefoot condition. The sagittal kinematic adaptations between conditions were found in the same way for all subjects and at the three running velocities. However, large individual variations were observed between the runners for the rearfoot kinematics.     

Tibiofemoral kinematics and condylar motion during the stance phase of gait

Accurate knowledge of the dynamic knee motion in-vivo is instrumental for understanding normal and pathological function of the knee joint. However, interpreting motion of the knee joint during gait in other than the sagittal plane remains controversial. In this study, we utilized the dual fluoroscopic imaging technique to investigate the six-degree-of-freedom kinematics and condylar motion of the knee during the stance phase of treadmill gait in eight healthy volunteers at a speed of 0.67 m/s. We hypothesized that the 6DOF knee kinematics measured during gait will be different from those reported for non-weightbearing activities, especially with regards to the phenomenon of femoral rollback. In addition, we hypothesized that motion of the medial femoral condyle in the transverse plane is greater than that of the lateral femoral condyle during the stance phase of treadmill gait. The rotational motion and the anterior–posterior translation of the femur with respect to the tibia showed a clear relationship with the flexion–extension path of the knee during the stance phase. Additionally, we observed that the phenomenon of femoral rollback was reversed, with the femur noted to move posteriorly with extension and anteriorly with flexion. Furthermore, we noted that motion of the medial femoral condyle in the transverse plane was greater than that of the lateral femoral condyle during the stance phase of gait (17.4±2.0 mm vs. 7.4±6.1 mm, respectively; p<0.01). The trend was opposite to what has been observed during non-weightbearing flexion or single-leg lunge in previous studies. These data provide baseline knowledge for the understanding of normal physiology and for the analysis of pathological function of the knee joint during walking. These findings further demonstrate that knee kinematics is activity-dependent and motion patterns of one activity (non-weightbearing flexion or lunge) cannot be generalized to interpret a different one (gait).     

Cross-correlation between EMG and center of gravity during quiet stance: theory and simulations

Several signal processing tools have been employed in the experimental study of the postural control system in humans. Among them, the cross-correlation function has been used to analyze the time relationship between signals such as the electromyogram and the horizontal projection of the center of gravity. The common finding is that the electromyogram precedes the biomechanical signal, a result that has been interpreted in different ways, for example, the existence of feedforward control or the preponderance of a velocity feedback. It is shown here, analytically and by simulation, that the cross-correlation function is depedent in a complicated way on system parameters and on noise spectra. Results similar to those found experimentally, e.g., electromyiogram preceding the biomechanical signal may be obtained in a postural control model without any feedforward control and without any velocity feedback. Therefore, correct interpretations of experimentally obtained cross-correlation functions may require additional information about the system. The results extend to other biomedical applications where two signals from a closed loop system are cross-correlated.     

Direct corticospinal pathways contribute to neuromuscular control of perturbed stance.

The antigravity soleus muscle (Sol) is crucial for compensation of stance perturbation. A corticospinal contribution to the compensatory response of the Sol is under debate. The present study assessed spinal, corticospinal, and cortical excitability at the peaks of short- (SLR), medium- (MLR), and long-latency responses (LLR) after posterior translation of the feet. Transcranial magnetic stimulation (TMS) and peripheral nerve stimulation were individually adjusted so that the peaks of either motor evoked potential (MEP) or H reflex coincided with peaks of SLR, MLR, and LLR, respectively. The influence of specific, presumably direct, corticospinal pathways was investigated by H-reflex conditioning. When TMS was triggered so that the MEP arrived in the Sol at the same time as the peaks of SLR and MLR, EMG remained unaffected. Enhanced EMG was observed when the MEP coincided with the LLR peak (P < 0.001). Similarly, conditioning of the H reflex by subthreshold TMS facilitated H reflexes only at LLR (P < 0.001). The earliest facilitation after perturbation occurred after 86 ms. The TMS-induced H-reflex facilitation at LLR suggests that increased cortical excitability contributes to the augmentation of the LLR peaks. This provides evidence that the LLR in the Sol muscle is at least partly transcortical, involving direct corticospinal pathways. Additionally, these results demonstrate that approximately 86 ms after perturbation, postural compensatory responses are cortically mediated.     

Ankle torque control that shifts the center of pressure from heel to toe contributes non-zero sagittal plane angular momentum during human walking

A principle objective of human walking is controlling angular motion of the body as a whole to remain upright. The force of the ground on each foot (F) reflects that control, and recent studies show that in the sagittal plane F exhibits a specific coordination between F direction and center-of-pressure (CP) that is conducive to remaining upright. Typical walking involves the CP shifting relative to the body due to two factors: posterior motion of the foot with respect to the hip (stepping) and motion of the CP relative to the foot (foot roll-over). Recent research has also shown how adjusting ankle torque alone to shift CP relative to the foot systematically alters the direction of F, and thus, could play a key role in upright posture and the F measured during walking. This study explores how the CP shifts due to stepping and foot roll-over contribute to the observed F and its role in maintaining upright posture. Experimental walking kinetics and kinematics were combined with a mechanical model of the human to show that variation in F that was not attributable to foot roll-over had systematic correlation between direction and CP that could be described by an intersection point located near the center-of-mass. The findings characterize a component of walking motor control, describe how typical foot roll-over contributes to postural control, and provide a rationale for the increased fall risk observed in individuals with atypical ankle muscle function.