Dominic James Farris

The mechanics and energetics of human walking and running: a joint level perspective

Humans walk and run at a range of speeds. While steady locomotion at a given speed requires no net mechanical work, moving faster does demand both more positive and negative mechanical work per stride. Is this increased demand met by increasing power output at all lower limb joints or just some of them? Does running rely on different joints for power output than walking? How does this contribute to the metabolic cost of locomotion? This study examined the effects of walking and running speed on lower limb joint mechanics and metabolic cost of transport in humans. Kinematic and kinetic data for 10 participants were collected for a range of walking (0.75, 1.25, 1.75, 2.0 m s−1) and running (2.0, 2.25, 2.75, 3.25 m s−1) speeds. Net metabolic power was measured by indirect calorimetry. Within each gait, there was no difference in the proportion of power contributed by each joint (hip, knee, ankle) to total power across speeds. Changing from walking to running resulted in a significant (p = 0.02) shift in power production from the hip to the ankle which may explain the higher efficiency of running at speeds above 2.0 m s−1 and shed light on a potential mechanism behind the walk–run transition.

Human medial gastrocnemius force–velocity behavior shifts with locomotion speed and gait

Humans walk and run over a wide range of speeds with remarkable efficiency. For steady locomotion, moving at different speeds requires the muscle–tendon units of the leg to modulate the amount of mechanical power the limb absorbs and outputs in each step. How individual muscles adapt their behavior to modulate limb power output has been examined using computer simulation and animal models, but has not been studied in vivo in humans. In this study, we used a combination of ultrasound imaging and motion analysis to examine how medial gastrocnemius (MG) muscle–tendon unit behavior is adjusted to meet the varying mechanical demands of different locomotor speeds during walking and running in humans. The results highlighted key differences in MG fascicle-shortening velocity with both locomotor speed and gait. Fascicle-shortening velocity at the time of peak muscle force production increased with walking speed, impairing the ability of the muscle to produce high peak forces. Switching to a running gait at 2.0 m⋅s−1 caused fascicle shortening at the time of peak force production to shift to much slower velocities. This velocity shift facilitated a large increase in peak muscle force and an increase in MG power output. MG fascicle velocity may be a key factor that limits the speeds humans choose to walk at, and may explain the transition from walking to running. This finding is consistent with previous modeling studies.

Could intra-tendinous hyperthermia during running explain chronic injury of the human Achilles tendon?

Chronic tendinopathy of the human Achilles tendon (AT) is common but its injury mechanism is not fully understood. It has been hypothesised that heat energy losses from the AT during running could explain the degeneration of AT material seen with injury. A mathematical model of AT temperature distribution was used to predict what temperatures the core of the AT could reach during running. This model required input values for mechanical properties of the AT (stiffness, hysteresis, cross-sectional area (CSA), strain during running) which were determined using a combination of ultrasound imaging, kinematic and kinetic data. AT length data were obtained during hopping and treadmill running (12 kmph) using ultrasound images of the medial gastrocnemius (50 Hz) and kinematic data (200 Hz). AT force data were calculated from inverse dynamics during hopping and combined with AT length data to compute AT stiffness and hysteresis. AT strain was computed from AT length data during treadmill running. AT CSA was measured on transverse ultrasound scans of the AT. Mean ± sd tendon properties were: stiffness = 176 ± 41 Nmm(-1), hysteresis =17 ± 12%, strain during running =3.5 ± 1.8% and CSA = 42 ± 8 mm(2). These values were input into the model of AT core temperature and this was predicted to reach at least 41°C during running. Such temperatures were deemed to be conservative estimates but still sufficient for tendon hyperthermia to be a potential cause of tendon injury.

Linking the mechanics and energetics of hopping with elastic ankle exoskeletons

The springlike mechanics of the human leg during bouncing gaits has inspired the design of passive assistive devices that use springs to aid locomotion. The purpose of this study was to test whether a passive spring-loaded ankle exoskeleton could reduce the mechanical and energetic demands of bilateral hopping on the musculoskeletal system. Joint level kinematics and kinetics were collected with electromyographic and metabolic energy consumption data for seven participants hopping at four frequencies (2.2, 2.5, 2.8, and 3.2 Hz). Hopping was performed without an exoskeleton; with an springless exoskeleton; and with a spring-loaded exoskeleton. Spring-loaded ankle exoskeletons reduced plantar flexor muscle activity and the biological contribution to ankle joint moment (15-25%) and average positive power (20-40%). They also facilitated reductions in metabolic power (15-20%) across frequencies from 2.2 to 2.8 Hz compared with hopping with a springless exoskeleton. Reductions in metabolic power compared with hopping with no exoskeleton were restricted to hopping at 2.5 Hz only (12%). These results highlighted the importance of reducing the rate of muscular force production and work to achieve metabolic reductions. They also highlighted the importance of assisting muscles acting at the knee joint. Exoskeleton designs may need to be tuned to optimize exoskeleton mass, spring stiffness, and spring slack length to achieve greater metabolic reductions.

Elastic ankle exoskeletons reduce soleus muscle force but not work in human hopping

Inspired by elastic energy storage and return in tendons of human leg muscle-tendon units (MTU), exoskeletons often place a spring in parallel with an MTU to assist the MTU. However, this might perturb the normally efficient MTU mechanics and actually increase active muscle mechanical work. This study tested the effects of elastic parallel assistance on MTU mechanics. Participants hopped with and without spring-loaded ankle exoskeletons that assisted plantar flexion. An inverse dynamics analysis, combined with in vivo ultrasound imaging of soleus fascicles and surface electromyography, was used to determine muscle-tendon mechanics and activations. Whole body net metabolic power was obtained from indirect calorimetry. When hopping with spring-loaded exoskeletons, soleus activation was reduced (30-70%) and so was the magnitude of soleus force (peak force reduced by 30%) and the average rate of soleus force generation (by 50%). Although forces were lower, average positive fascicle power remained unchanged, owing to increased fascicle excursion (+4-5 mm). Net metabolic power was reduced with exoskeleton assistance (19%). These findings highlighted that parallel assistance to a muscle with appreciable series elasticity may have some negative consequences, and that the metabolic cost associated with generating force may be more pronounced than the cost of doing work for these muscles.

Differential strain patterns of the human Achilles tendon determined in vivo with freehand three-dimensional ultrasound imaging

SUMMARY The human Achilles tendon (AT) has often been considered to act as a single elastic structure in series with the muscles of the triceps surae. As such it has been commonly modelled as a Hookean spring of uniform stiffness. However, the free AT and the proximal AT have distinctly different structures that lend themselves to different elastic properties. This study aimed to use three-dimensional freehand ultrasound imaging to determine whether the proximal AT and the free AT exhibit different elastic behaviour during sub-maximal, fixed-end contractions of the triceps surae. Six male and five female participants (mean ± s.d. age=27±5 years) performed fixed position contractions of the plantar-flexors on an isokinetic dynamometer at 50% of their maximum voluntary contraction in this position. Freehand three-dimensional ultrasound imaging was used to reconstruct the free-tendon and proximal AT at rest and during contraction. The free-tendon exhibited significantly (P=0.03) greater longitudinal strain (5.2±1.7%) than the proximal AT (2.6±2.0%). The lesser longitudinal strain of the proximal AT was linked to the fact that it exhibited considerable transverse (orthogonal to the longitudinal direction) strains (5.0±4%). The transverse strain of the proximal AT is likely due to the triceps surae muscles bulging upon contraction, and thus the level of bulging may influence the elastic behaviour of the proximal AT. This might have implications for the understanding of triceps surae muscle–tendon interaction during locomotion, tendon injury mechanics and previous measurements of AT elastic properties.

Musculoskeletal modelling deconstructs the paradoxical effects of elastic ankle exoskeletons on plantar-flexor mechanics and energetics during hopping.

Experiments have shown that elastic ankle exoskeletons can be used to reduce ankle joint and plantar-flexor muscle loading when hopping in place and, in turn, reduce metabolic energy consumption. However, recent experimental work has shown that such exoskeletons cause less favourable soleus (SO) muscle–tendon mechanics than is observed during normal hopping, which might limit the capacity of the exoskeleton to reduce energy consumption. To directly link plantar-flexor mechanics and energy consumption when hopping in exoskeletons, we used a musculoskeletal model of the human leg and a model of muscle energetics in simulations of muscle–tendon dynamics during hopping with and without elastic ankle exoskeletons. Simulations were driven by experimental electromyograms, joint kinematics and exoskeleton torque taken from previously published data. The data were from seven males who hopped at 2.5 Hz with and without elastic ankle exoskeletons. The energetics model showed that the total rate of metabolic energy consumption by ankle muscles was not significantly reduced by an ankle exoskeleton. This was despite large reductions in plantar-flexor force production (40–50%). The lack of larger metabolic reductions with exoskeletons was attributed to increases in plantar-flexor muscle fibre velocities and a shift to less favourable muscle fibre lengths during active force production. This limited the capacity for plantar-flexors to reduce activation and energy consumption when hopping with exoskeleton assistance.

The role of human ankle plantar flexor muscle-tendon interaction and architecture in maximal vertical jumping examined in vivo.

ABSTRACT Humans utilise elastic tendons of lower limb muscles to store and return energy during walking, running and jumping. Anuran and insect species use skeletal structures and/or dynamics in conjunction with similarly compliant structures to amplify muscle power output during jumping. We sought to examine whether human jumpers use similar mechanisms to aid elastic energy usage in the plantar flexor muscles during maximal vertical jumping. Ten male athletes performed maximal vertical squat jumps. Three-dimensional motion capture and a musculoskeletal model were used to determine lower limb kinematics that were combined with ground reaction force data in an inverse dynamics analysis. B-mode ultrasound imaging of the lateral gastrocnemius (GAS) and soleus (SOL) muscles was used to measure muscle fascicle lengths and pennation angles during jumping. Our results highlighted that both GAS and SOL utilised stretch and recoil of their series elastic elements (SEEs) in a catapult-like fashion, which likely serves to maximise ankle joint power. The resistance of supporting of body weight allowed initial stretch of both GAS and SOL SEEs. A proximal-to-distal sequence of joint moments and decreasing effective mechanical advantage early in the extension phase of the jumping movement were observed. This facilitated a further stretch of the SEE of the biarticular GAS and delayed recoil of the SOL SEE. However, effective mechanical advantage did not increase late in the jump to aid recoil of elastic tissues. Summary: Elastic power delivered from ankle muscle-tendons during jumping in humans is aided by energy storage against body weight and proximal-distal sequencing of limb action, but not variable mechanical advantage, which may explain our modest jumping ability.

Shoes alter the spring-like function of the human foot during running

The capacity to store and return energy in legs and feet that behave like springs is crucial to human running economy. Recent comparisons of shod and barefoot running have led to suggestions that modern running shoes may actually impede leg and foot-spring function by reducing the contributions from the leg and foot musculature. Here we examined the effect of running shoes on foot longitudinal arch (LA) motion and activation of the intrinsic foot muscles. Participants ran on a force-instrumented treadmill with and without running shoes. We recorded foot kinematics and muscle activation of the intrinsic foot muscles using intramuscular electromyography. In contrast to previous assertions, we observed an increase in both the peak (flexor digitorum brevis +60%) and total stance muscle activation (flexor digitorum brevis +70% and abductor hallucis +53%) of the intrinsic foot muscles when running with shoes. Increased intrinsic muscle activation corresponded with a reduction in LA compression (−25%). We confirm that running shoes do indeed influence the mechanical function of the foot. However, our findings suggest that these mechanical adjustments are likely to have occurred as a result of increased neuromuscular output, rather than impaired control as previously speculated. We propose a theoretical model for foot–shoe interaction to explain these novel findings.

Individual limb mechanical analysis of gait following stroke

The step-to-step transition of walking requires significant mechanical and metabolic energy to redirect the center of mass. Inter-limb mechanical asymmetries during the step-to-step transition may increase overall energy demands and require compensation during single-support. The purpose of this study was to compare individual limb mechanical gait asymmetries during the step-to-step transitions, single-support and over a complete stride between two groups of individuals following stroke stratified by gait speed (≥0.8 m/s or <0.8 m/s). Twenty-six individuals with chronic stroke walked on an instrumented treadmill to collect ground reaction force data. Using the individual limbs method, mechanical power produced on the center of mass was calculated during the trailing double-support, leading double-support, and single-support phases of a stride, as well as over a complete stride. Robust inter-limb asymmetries in mechanical power existed during walking after stroke; for both groups, the non-paretic limb produced significantly more positive net mechanical power than the paretic limb during all phases of a stride and over a complete stride. Interestingly, no differences in inter-limb mechanical power asymmetry were noted between groups based on walking speed, during any phase or over a complete stride. Paretic propulsion, however, was different between speed-based groups. The fact that paretic propulsion (calculated from anterior-posterior forces) is different between groups, but our measure of mechanical work (calculated from all three directions) is not, suggests that limb power output may be dominated by vertical components, which are required for upright support.