Ankle extensor proprioceptors contribute to the enhancement of the soleus EMG during the stance phase of human walking

A rapid plantar flexion perturbation applied to the ankle during the stance phase of the step cycle during human walking unloads the ankle extensors and produces a marked decline in the soleus EMG. This demonstrates that sensory activity contributes importantly to the enhancement of the ankle extensor muscle activation during human walking. On average, the EMG begins to decline approximately 52 ms after the perturbation. In contrast, a rapid dorsi flex ion perturbation produces a group Ia mediated short-latency stretch reflex burst with an onset latency of approximately 36 ms. The transmission of sensory traffic from the foot and ankle was suppressed in 10 subjects by an anaesthetic nerve block produced with local injections of lidocaine hydrochloride. The anaesthetic block had no effect on the stance phase soleus EMG, the latencies of the EMG responses, or the magnitude of the EMG decline following the plantar flexion perturbation. Therefore, it is more likely that proprioceptive afferents, rather than cutaneous afferents, contribute to the background soleus EMG during the late stance phase of the step cycle. The large difference in onset latencies between the short-latency reflex and unload responses suggests that the largest of the active group Ia afferents might not contribute strongly to the background soleus EMG, although it remains to be determined which of the proprioceptive pathways provide the more important contributions.     

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.     

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.     

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.     

The human ankle during walking: implications for design of biomimetic ankle prostheses

The non-disabled human ankle joint was examined during walking in an attempt to determine overall system characteristics for use in the design of ankle prostheses. The hypothesis of the study was that the quasi-stiffness of the ankle changes when walking at different walking speeds. The hypothesis was examined using sagittal plane ankle moment versus ankle angle curves from 24 able-bodied subjects walking over a range of speeds. The slopes of the moment versus ankle angle curves (quasi-stiffness) during loading appeared to change as speed was increased and the relationship between the moment and angle during loading became increasingly non-linear. The loading and unloading portions of the moment versus angle curves showed clockwise loops (hysteresis) at self-selected slow speeds that reduced essentially to zero as the speed increased to self-selected normal speeds. Above self-selected normal speeds, the loops started to traverse a counter-clockwise path that increased in area as the speed was increased. These characteristics imply that the human ankle joint could be effectively replaced with a rotational spring and damper for slow to normal walking speeds. However, to mimic the characteristics of the human ankle during walking at fast speeds, an augmented system would be necessary. This notion is supported by the sign of the ankle power at the time of opposite heel contact, which was negative for slow speeds, was near zero at normal speeds, and was positive for fast walking speeds.     

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).     

Estimation of passive ankle joint moment during standing and walking

This study estimated the passive ankle joint moment during standing and walking initiation and its contribution to total ankle joint moment during that time. The decrement of passive joint moment due to muscle fascicle shortening upon contraction was taken into account. Muscle fascicle length in the medial gastrocnemius, which was assumed to represent muscle fascicle length in plantarflexors, was measured using ultrasonography during standing, walking initiation, and cyclical slow passive ankle joint motion. Total ankle joint moment during standing and walking initiation was calculated from ground reaction forces and joint kinematics. Passive ankle joint moment during the cyclical ankle joint motion was measured via a dynamometer. Passive ankle joint moment during standing and at the time (Tp) when the MG muscle-tendon complex length was longest in the stance phase during walking initiation were 2.3 and 5.4 Nm, respectively. The muscle fascicle shortened by 2.9 mm during standing compared with the length at rest, which decreased the contribution of passive joint moment from 19.9% to 17.4%. The muscle fascicle shortened by 4.3 mm at Tp compared with the length at rest, which decreased the contribution of passive joint moment from 8.0% to 5.8%. These findings suggest that (a) passive ankle joint moment plays an important role during standing and walking initiation even in view of the decrement of passive joint moment due to muscle fascicle shortening upon muscle contraction, and (b) muscle fascicle shortening upon muscle contraction must be taken into account when estimating passive joint moment during movements.     

The natural frequency of the foot-surface cushion during the stance phase of running

Researchers have reported on the stiffness of running in holistic terms, i.e. for the structures that are undergoing deformation as a whole rather than in terms of specific locations. This study aimed to estimate both the natural frequency and the viscous damping coefficient of the human foot-surface cushion, during the period between the heel strike and the mid-stance phase of running, using a purposely developed one degree-of-freedom inverted pendulum state space model of the leg. The model, which was validated via a comparison of measured and estimated ground reaction forces, incorporated a novel use of linearized and extended Kalman filter estimators. Investigation of the effect of variation of the natural frequency and/or the damping of the cushioning mechanism during running, using the said model, revealed the natural frequency of running on said foot-surface cushion, during the stance phase, to lie between 5 and 11 Hz. The "extended Kalman filter (EKF)" approach, that was used here for the first time to directly apply measured ground forces, may be widely applicable to the identification process of combined estimation of both unknown physiological state and mechanical characteristics of the environment in an inverse dynamic model.     

Bone contact forces on the distal tibia during the stance phase of running

Although the tibia is a common site of stress fractures in runners, the loading of the tibia during running is not well understood. An integrated experimental and modeling approach was therefore used to estimate the bone contact forces acting on the distal end of the tibia during the stance phase of running, and the contributions of external and internal sources to these forces. Motion capture and force plate data were recorded for 10 male runners as they ran at 3.5-4 m/s. From these data, the joint reaction force (JRF), muscle forces, and bone contact force on the tibia were computed at the ankle using inverse dynamics and optimization methods. The distal end of the tibia was compressed and sheared posteriorly throughout most of stance, with respective peak forces of 9.00+/-1.13 and 0.57+/-0.18 body weights occurring during mid stance. Internal muscle forces were the primary source of tibial compression, whereas the JRF was the primary source of tibial shear due to the forward inclination of the leg relative to the external ground reaction force. The muscle forces and JRF both acted to compress the tibia, but induced tibial shear forces in opposing directions during stance, magnifying tibial compression and reducing tibial shear. The superposition of the peak compressive and posterior shear forces at mid stance may contribute to stress fractures in the posterior face of the tibia. The implications are that changes in running technique could potentially reduce stress fracture risk.     

The role of phase shifts of sensory inputs in walking revealed by means of phase reduction

Detailed neural network models of animal locomotion are important means to understand the underlying mechanisms that control the coordinated movement of individual limbs. Daun-Gruhn and Tóth, Journal of Computational Neuroscience 31(2), 43–60 (2011) constructed an inter-segmental network model of stick insect locomotion consisting of three interconnected central pattern generators (CPGs) that are associated with the protraction-retraction movements of the front, middle and hind leg. This model could reproduce the basic locomotion coordination patterns, such as tri- and tetrapod, and the transitions between them. However, the analysis of such a system is a formidable task because of its large number of variables and parameters. In this study, we employed phase reduction and averaging theory to this large network model in order to reduce it to a system of coupled phase oscillators. This enabled us to analyze the complex behavior of the system in a reduced parameter space. In this paper, we show that the reduced model reproduces the results of the original model. By analyzing the interaction of just two coupled phase oscillators, we found that the neighboring CPGs could operate within distinct regimes, depending on the phase shift between the sensory inputs from the extremities and the phases of the individual CPGs. We demonstrate that this dependence is essential to produce different coordination patterns and the transition between them. Additionally, applying averaging theory to the system of all three phase oscillators, we calculate the stable fixed points - they correspond to stable tripod or tetrapod coordination patterns and identify two ways of transition between them.     

Possible functional roles of phase resetting during walking

The walking rhythm is known to show phase shift or "reset" in response to external impulsive perturbations. We tried to elucidate functional roles of the phase reset possibly used for the neural control of locomotion. To this end, a system with a double pendulum as a simplified model of the locomotor control and a model of bipedal locomotion were employed and analyzed in detail. In these models, a movement corresponding to the normal steady-state walking was realized as a stable limit cycle solution of the system. Unexpected external perturbations applied to the system can push the state point of the system away from its limit cycle, either outside or inside the basin of attraction of the limit cycle. Our mathematical analyses of the models suggested functional roles of the phase reset during walking as follows. Function 1: an appropriate amount of the phase reset for a given perturbation can contribute to relocating the system's state point outside the basin of attraction of the limit cycle back to the inside. Function 2: it can also be useful to reduce the convergence time (the time necessary for the state point to return to the limit cycle). In experimental studies during walking of animals and humans, the reset of walking rhythm induced by perturbations was investigated using the phase transition curve (PTC) or the phase resetting curve (PRC) representing phase-dependent responses of the walking. We showed, for the simple double-pendulum model, the existence of the optimal phase control and the corresponding PTC that could optimally realize the aforementioned functions in response to impulsive force perturbations. Moreover, possible forms of PRC that can avoid falling against the force perturbations were predicted by the biped model, and they were compared with the experimentally observed PRC during human walking. Finally, physiological implications of the results were discussed.     

Adaptive feedback potential in dynamic stability during disturbed walking in the elderly

After perturbation of the gait, feedback information may help regaining balance adequately, but it remains unknown whether adaptive feedback responses are possible after repetitive and unexpected perturbations during gait and if there are age-related differences. Prior experience may contribute to improved reactive behavior. Fourteen old (59-73 yrs) and fourteen young (22-31 yrs) males walked on a walkway which included one covered element. By exchanging this element participants either stepped on hard surface or unexpectedly on soft surface which caused a perturbation in gait. The gait protocol contained 5 unexpected soft trials to quantify the reactive adaptation. Each soft trial was followed by 4-8 hard trials to generate a wash-out effect. The dynamic stability was investigated by using the margin of stability (MoS), which was calculated as the difference between the anterior boundary of the base of support and the extrapolated position of the center of mass in the anterior-posterior direction. MoS at recovery leg touchdown were significantly lower in the unexpected soft trials compared to the baseline, indicating a less stable posture. However, MoS increased (p<0.05) in both groups within the disturbed trials, indicating feedback adaptive improvements. Young and old participants showed differences in the handling of the perturbation in the course of several trials. The magnitude of the reactive adaptation after the fifth unexpected perturbation was significantly different compared to the first unexpected perturbation (old: 49±30%; young: 77±40%), showing a tendency (p=0.065) for higher values in the young participants. Old individuals maintain the ability to adapt to feedback controlled perturbations. However, the locomotor behavior is more conservative compared to the young ones, leading to disadvantages in the reactive adaptation during disturbed walking.     

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

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

Contribution to histochemical distribution of non-specific cholinesterase activity in sensory ganglia of some mammals

The activity of non-specific cholinesterase was demonstrated histochemically in satellite cells of the spinal ganglia from adult rat, cat, rabbit and baboon. The spinal ganglia of newborn rats displayed distinct intraneuronal reactivity for non-specific cholinesterase while a low reactivity was observed in satellite cells. The spinal and trigeminal ganglia of adult mice contained satellite cells with non-specific cholinesterase reactivity only sporadically. Most of reaction product for non-specific cholinesterase activity (from low to high intensity) was found in perikarya of the neurons. Spinal and trigeminal ganglia of the same mice embryo exhibited diffuse staining for non-specific cholinesterase activity remaining in the spinal ganglia of newborn mice. The trigeminal ganglia of newborn mice exhibited, however, more differentiated pattern of the positive reaction for non-specific cholinesterase like adult animals. The pattern of histochemical distribution of non-specific cholinesterase activity in trigeminal and spinal ganglia from mice of various ages corresponds with morphological differentiation and maturation undergoing in a rostrocaudal wave. Intraneuronal presence of non-specific cholinesterase activity in sensory ganglia during development and in adult animals gives a new possibilities for explanation of the functional involvement of this enzyme in the nervous system.     

Contributions of the individual ankle plantar flexors to support, forward progression and swing initiation during walking.

Walking is a motor task requiring coordination of many muscles. Previous biomechanical studies, based primarily on analyses of the net ankle moment during stance, have concluded different functional roles for the plantar flexors. We hypothesize that some of the disparities in interpretation arise because of the effects of the uniarticular and biarticular muscles that comprise the plantar flexor group have not been separated. Furthermore, we believe that an accurate determination of muscle function requires quantification of the contributions of individual plantar flexor muscles to the energetics of individual body segments. In this study, we examined the individual contributions of the ankle plantar flexors (gastrocnemius (GAS); soleus (SOL)) to the body segment energetics using a musculoskeletal model and optimization framework to generate a forward dynamics simulation of normal walking at 1.5 m/s. At any instant in the gait cycle, the contribution of a muscle to support and forward progression was defined by its contribution to trunk vertical and horizontal acceleration, respectively, and its contribution to swing initiation by the mechanical energy it delivers to the leg in pre-swing (i.e., double-leg stance prior to toe-off). GAS and SOL were both found to provide trunk support during single-leg stance and pre-swing. In early single-leg stance, undergoing eccentric and isometric activity, they accelerate the trunk vertically but decelerate forward trunk progression. In mid single-leg stance, while isometric, GAS delivers energy to the leg while SOL decelerates it, and SOL delivers energy to the trunk while GAS decelerates it. In late single-leg stance through pre-swing, though GAS and SOL both undergo concentric activity and accelerate the trunk forward while decelerating the downward motion of the trunk (i.e., providing forward progression and support), they execute different energetic functions. The energy produced from SOL accelerates the trunk forward, whereas GAS delivers almost all its energy to accelerate the leg to initiate swing. Although GAS and SOL maintain or accelerate forward motion in mid single-leg stance through pre-swing, other muscles acting at the beginning of stance contribute comparably to forward progression. In summary, throughout single-leg stance both SOL and GAS provide vertical support, in mid single-leg stance SOL and GAS have opposite energetic effects on the leg and trunk to ensure support and forward progression of both the leg and trunk, and in pre-swing only GAS contributes to swing initiation.     

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.