SUMMARYThe musculoskeletal structure of the foot and ankle has the potential to influence human sprinting performance in complex ways. A large Achillesʼ tendon moment arm improves the mechanical advantage of the triceps surae but also produces larger shortening velocity during rapid plantarflexion, which detracts from the force-generating capacity of the plantarflexors. The lever arm of the ground reaction force that resists the muscular plantarflexor moment during propulsive push-off is constrained in part by the skeletal structure of the foot. In this study, we measured the plantarflexion moment arms of the Achillesʼ tendon, lateral gastrocnemius fascicle lengths and pennation angles, and anthropometric characteristics of the foot and lower leg in collegiate sprinters and height-matched non-sprinters. The Achillesʼ tendon moment arms of the sprinters were 25% smaller on average in sprinters than in non-sprinters (P<0.001) whereas the sprintersʼ fascicles were 11% longer on average (P0.024). The ratio of fascicle length to moment arm was 50% larger in sprinters (P<0.001). Sprinters were found to have longer toes (P0.032) and shorter lower legs (P0.026) than non sprinters. A simple computer simulation of the sprint push-off demonstrated that shorter plantarflexor moment arms and longer toes, like those measured in sprinters, permit greater generation of forward impulse. Simulated propulsion was enhanced in both cases by increasing the ʻgear ratioʼ of the foot, thus maintaining plantarflexor fibre length and reducing peak fibre shortening velocity. Longer toes especially prolonged the time of contact, giving greater time for forward acceleration by propulsive ground reaction force.

During muscle contractions, the muscle fascicles may shorten at a rate different from the muscletendon unit, and the ratio of these velocities is its gearing. Appropriate gearing allows fascicles to reduce their shortening velocities and allows them to operate at effective shortening velocities across a range of movements. Gearing of the muscle fascicles within the muscle belly is the result of rotations of the fascicles and bulging of the belly. Variable gearing can also occur as a result of tendon length changes that can be caused by changes in the relative timing of muscle activity for different mechanical tasks. Recruitment patterns of slow and fast fibres are crucial for achieving optimal muscle performance, and coordination between muscles is related to whole limb performance. Poor coordination leads to inefficiencies and loss of power, and optimal coordination is required for high power outputs and high mechanical efficiencies from the limb. This paper summarizes key studies in these areas of neuromuscular mechanics and results from studies where we have tested these phenomena on a cycle ergometer are presented to highlight novel insights. The studies show how muscle structure and neural activation interact to generate smooth and effective motion of the body.Keywords: power; efficiency; gearing; pennation PROLOGUEMuscles have a surprising variety of functions during locomotion and can serve as motors, brakes, springs and struts [1]. These muscle functions depend on the arrangement of the muscle fibres within the muscle belly, the length and velocity of the fibres and also on the timing of their activation relative to those length changes. When the muscle fibres undergo substantial length change the muscle can act as a motor if it is active during shortening, or a brake if it is active during lengthening; when the length change is minimal then the muscle can act like a strut if it actively develops force to resist stretch [1]. Muscle fibre velocity may differ from the velocity of the muscletendon unit (MTU) in a process known as gearing. Thus, the extent to which the muscle fibres change length during a limb motion depends on how they are geared relative to the MTU. The timing and the level of activation are under neural control. This paper is a synthesis of previous literature on both gearing and activation, with a particular focus on some of the more recent findings in man. We introduce additional findings about structural mechanics in man and propose that some of the features of muscle gearing actually result from the interactions between the structural mechanics and the activation patterns of the muscles. MUSCLE STRUCTURE AND FASCICLE GEARINGMTUs are comprised of both tendinous structures containing a large proportion of collagen fibres, and skeletal muscle fibres containing the actin-and myosin-rich contractile machinery. The skeletal muscle fibres typically lie parallel to each other and are bundled to form fascicles that combine to form the muscle belly. The mechanics of fibres and fascicles can be ...

The following papers by Richard Lieber (Skeletal Muscle as an Actuator), Thomas Roberts (Elastic Mechanisms and Muscle Function), Silvia Blemker (Skeletal Muscle has a Mind of its Own: a Computational Framework to Model the Complex Process of Muscle Adaptation) and Sabrina Lee (Muscle Properties of Spastic Muscle (Stroke and CP) are summaries of their representative contributions for the session on skeletal muscle mechanics, energetics and plasticity at the 2016 Biomechanics and Neural Control of Movement Conference (BANCOM 2016). Dr. Lieber revisits the topic of sarcomere length as a fundamental property of skeletal muscle contraction. Specifically, problems associated with sarcomere length non-uniformity and the role of sarcomerogenesis in diseases such as cerebral palsy are critically discussed. Dr. Roberts then makes us aware of the (often neglected) role of the passive tissues in muscles and discusses the properties of parallel elasticity and series elasticity, and their role in muscle function. Specifically, he identifies the merits of analyzing muscle deformations in three dimensions (rather than just two), because of the potential decoupling of the parallel elastic element length from the contractile element length, and reviews the associated implications for the architectural gear ratio of skeletal muscle contraction. Dr. Blemker then tackles muscle adaptation using a novel way of looking at adaptive processes and what might drive adaptation. She argues that cells do not have pre-programmed behaviors that are controlled by the nervous system. Rather, the adaptive responses of muscle fibers are determined by sub-cellular signaling pathways that are affected by mechanical and biochemical stimuli; an exciting framework with lots of potential. Finally, Dr. Lee takes on the challenging task of determining human muscle properties in vivo. She identifies the dilemma of how we can demonstrate the effectiveness of a treatment, specifically in cases of muscle spasticity following stroke or in children with cerebral palsy. She then discusses the merits of ultrasound based elastography, and the clinical possibilities this technique might hold. Overall, we are treated to a vast array of basic and clinical problems in skeletal muscle mechanics and physiology, with some solutions, and many suggestions for future research.

Muscle stiffness and muscle composition, as indicated by SW speed and echo intensity, may be altered in stroke-impaired muscle at rest. These findings highlight the potential for SW elastography as a tool for both investigating the fundamental mechanisms behind changes in stroke-impaired muscle, and for evaluation of muscle mechanical properties as part of clinical examination.

Recruitment patterns and activation dynamics of different motor units greatly influence the temporal pattern and magnitude of muscle force development, yet these features are not often considered in muscle models. The purpose of this study was to characterize the recruitment and activation dynamics of slow and fast motor units from electromyographic (EMG) recordings and twitch force profiles recorded directly from animal muscles. EMG and force data from the gastrocnemius muscles of seven goats were recorded during in vivo tendon-tap reflex and in situ nerve stimulation experiments. These experiments elicited EMG signals with significant differences in frequency content (p<0.001). The frequency content was characterized using wavelet and principal components analysis, and optimized wavelets with centre frequencies, 149.94Hz and 323.13Hz, were obtained. The optimized wavelets were used to calculate the EMG intensities and, with the reconstructed twitch force profiles, to derive transfer functions for slow and fast motor units that estimate the activation state of the muscle from the EMG signal. The resulting activation-deactivation time constants gave r values of 0.98 to 0.99 between the activation state and the force profiles. This work establishes a framework for developing improved muscle models that consider the intrinsic properties of slow and fast fibres within a mixed muscle, and that can more accurately predict muscle force output from EMG.

Background Individuals with cerebral palsy tend to have altered muscle architecture and composition, but little is known about the muscle material properties, specifically stiffness. Shear wave ultrasound elastography allows shear wave speed, which is related to stiffness, to be measured in vivo in individual muscles. Our aim was to evaluate the material properties, specifically stiffness, as measured by shear wave speed of the medial gastrocnemius and tibialis anterior muscles in children with hemiplegic cerebral palsy across a range of ankle torques and positions, and fascicle strains. Method Shear wave speed was measured bilaterally in the medial gastrocnemius and tibialis anterior over a range of ankle positions and torques using shear wave ultrasound elastography in eight individuals with hemiplegic cerebral palsy. B-mode ultrasound was used to measure muscle thickness and fascicle strain. Results Shear waves traveled faster in the medial gastrocnemius and tibialis anterior of the more-affected limb by 14% (p=0.024) and 20% (p=0.03), respectively, when the ankle was at 90°. Shear wave speed in the medial gastrocnemius increased as the ankle moved from plantarflexion to dorsiflexion (less affected: r2=0.82,p<0.001; more-affected: r2=0.69,p<0.001) and as ankle torque increased (less affected: r2=0.56,p<0.001; more-affected: r2=0.45,p<0.001). In addition, shear wave speed was strongly correlated with fascicle strain (less affected: r2=0.63,p<0.001; more-affected: r2=0.53,p<0.001). Interpretation The higher shear wave speed in the more-affected limb of individuals with cerebral palsy indicates greater muscle stiffness, and demonstrates the clinical potential of shear wave elastography as a non-invasive tool for investigating mechanisms of altered muscle properties and informing diagnosis and treatment.

Biomechanical models of whole muscles commonly used in simulations of musculoskeletal function and movement typically assume that the muscle generates force as a scaled-up muscle fiber. However, muscles are comprised of motor units that have different intrinsic properties and that can be activated at different times. This study tested whether a muscle model comprised of motor units that could be independently activated resulted in more accurate predictions of force than traditional Hill-type models. Forces predicted by the models were evaluated by direct comparison with the muscle forces measured in situ from the gastrocnemii in goats. The muscle was stimulated tetanically at a range of frequencies, muscle fiber strains were measured using sonomicrometry, and the activation patterns of the different types of motor unit were calculated from electromyographic recordings. Activation patterns were input into five different muscle models. Four models were traditional Hill-type models with different intrinsic speeds and fiber-type properties. The fifth model incorporated differential groups of fast and slow motor units. For all goats, muscles and stimulation frequencies the differential model resulted in the best predictions of muscle force. The in situ muscle output was shown to depend on the recruitment of different motor units within the muscle.

Hill-type models are commonly used to estimate muscle forces during human and animal movement —yet the accuracy of the forces estimated during walking, running, and other tasks remains largely unknown. Further, most Hill-type models assume a single contractile element, despite evidence that faster and slower motor units, which have different activation-deactivation dynamics, may be independently or collectively excited. This study evaluated a novel, two-element Hill-type model with “differential” activation of fast and slow contractile elements. Model performance was assessed using a comprehensive data set (including measures of EMG intensity, fascicle length, and tendon force) collected from the gastrocnemius muscles of goats during locomotor experiments. Muscle forces predicted by the new two-element model were compared to the forces estimated using traditional one-element models and to the forces measured in vivo using tendon buckle transducers. Overall, the two-element model resulted in the best predictions of in vivo gastrocnemius force. The coefficient of determination, r2, was up to 26.9% higher and the root mean square error, RMSE, was up to 37.4% lower for the two-element model than for the one-element models tested. All models captured salient features of the measured muscle force during walking, trotting, and galloping (r2 = 0.26 to 0.51), and all exhibited some errors (RMSE = 9.63 to 32.2% of the maximum in vivo force). These comparisons provide important insight into the accuracy of Hill-type models. The results also show that incorporation of fast and slow contractile elements within muscle models can improve estimates of time-varying, whole muscle force during locomotor tasks.