Humans trade off whole-body energy cost to avoid overburdening muscles while walking.

Metabolic cost minimization is thought to underscore the neural control of locomotion. Yet, avoiding high muscle activation, a cause of fatigue, often outperforms energy minimization in computational predictions of human gait. Discerning the relative importance of these criteria in human walking has proved elusive, in part, because they have not been empirically decoupled. Here, we explicitly decouple whole-body metabolic cost and 'fatigue-like' muscle activation costs (estimated from electromyography) by pitting them against one another using two distinct gait tasks. When experiencing these competing costs, participants (n = 10) chose the task that avoided overburdening muscles (fatigue avoidance) at the expense of higher metabolic power (p < 0.05). Muscle volume-normalized activation more closely models energy use and was also minimized by the participants' decision (p < 0.05), demonstrating that muscle activation was, at best, an inaccurate signal for metabolic energy. Energy minimization was only observed when there was no adverse effect on muscle activation costs. By decoupling whole-body metabolic and muscle activation costs, we provide among the first empirical evidence of humans embracing non-energetic optimality in favour of a clearly defined neuromuscular objective. This finding indicates that local muscle fatigue and effort may well be key factors dictating human walking behaviour and its evolution.

Plasticity of the gastrocnemius elastic system in response to decreased work and power demand during growth.

Elastic energy storage and release can enhance performance that would otherwise be limited by the force-velocity constraints of muscle. Although functional influence of a biological spring depends on tuning between components of an elastic system (the muscle, spring-driven mass and lever system), we do not know whether elastic systems systematically adapt to functional demand. To test whether altering work and power generation during maturation alters the morphology of an elastic system, we prevented growing guinea fowl (Numida meleagris) from jumping. We compared the jump performance of our treatment group at maturity with that of controls and measured the morphology of the gastrocnemius elastic system. We found that restricted birds jumped with lower jump power and work, yet there were no significant between-group differences in the components of the elastic system. Further, subject-specific models revealed no difference in energy storage capacity between groups, though energy storage was most sensitive to variations in muscle properties (most significantly operating length and least dependent on tendon stiffness). We conclude that the gastrocnemius elastic system in the guinea fowl displays little to no plastic response to decreased demand during growth and hypothesize that neural plasticity may explain performance variation.

Is conservation of center of mass mechanics a priority in human walking? Insights from leg-length asymmetry experiments.

Center of mass (COM) control has been proposed to serve economy- and stability-related locomotor task objectives. However, given the lack of evidence supporting direct sensing and/or regulation of the COM, it remains unclear whether COM mechanics are prioritized in the control scheme of walking. We posit that peripheral musculoskeletal structures, e.g. muscle, are more realistic control targets than the COM, given their abundance of sensorimotor receptors and ability to influence whole-body energetics. As a first test of this hypothesis, we examined whether conservation of stance-phase joint mechanics is prioritized over COM mechanics in a locomotor task where simultaneous conservation of COM and joint mechanics is not feasible: imposed leg-length asymmetry. Positive joint mechanical cost of transport (work per distance traveled; COTJNT) was maintained at values closer to normal walking than COM mechanical cost of transport (COTCOM; P<0.05, N=15). Furthermore, compared with our measures of COM mechanics (COTCOM, COM displacement), joint-level variables (COTJNT, integrated total support moment) also displayed stronger conservation (less change from normal walking) when the participants' self-selected gait was assessed against other possible gait solutions. We conclude that when walking humans are exposed to an asymmetric leg-length perturbation, control of joint mechanics is prioritized over COM mechanics. Our results suggest that mechanical and metabolic effort is likely regulated via control of peripheral structures and not directly at the level of the COM. Joint mechanics may provide a more accurate representation of the underlying locomotor control targets and may prove advantageous in informing predictive models of human walking.

Modulation of joint and limb mechanical work in walk-to-run transition steps in humans.

Surprisingly little information exists of the mechanics in the steps initializing the walk-to-run transition (WRT) in humans. Here, we assess how mechanical work of the limbs (vertical and horizontal) and the individual joints (ankle, knee and hip) are modulated as humans transition from a preferred constant walking velocity (vwalk) to a variety of running velocities (vrun; ranging from a sprint to a velocity slower than vwalk). WRTs to fast vrun values occur nearly exclusively through positive horizontal limb work, satisfying the goal of forward acceleration. Contrary to our hypothesis, however, positive mechanical work remains above that at vwalk even when decelerating. In these WRTs to slow running, positive mechanical work is remarkably high and is comprised nearly exclusively of vertical limb work. Vertical-to-horizontal work modulation may represent an optimization for achieving minimal and maximal vrun, respectively, while fulfilling an apparent necessity for energy input when initiating WRTs. Net work of the WRT steps was more evenly distributed across the ankle, knee and hip joints than expected. Absolute positive mechanical work exhibited a clearer modulation towards hip-based work at high accelerations (>3 m s-2), corroborating previous suggestions that the most proximal joints are preferentially recruited for locomotor tasks requiring high power and work production. In WRTs to very slow vrun values, high positive work is nevertheless done at the knee, indicating that modulation of joint work is not only dependent on the amount of work required but also the locomotor context.

Adaptive Remodeling of Achilles Tendon: A Multi-scale Computational Model.

While it is known that musculotendon units adapt to their load environments, there is only a limited understanding of tendon adaptation in vivo. Here we develop a computational model of tendon remodeling based on the premise that mechanical damage and tenocyte-mediated tendon damage and repair processes modify the distribution of its collagen fiber lengths. We explain how these processes enable the tendon to geometrically adapt to its load conditions. Based on known biological processes, mechanical and strain-dependent proteolytic fiber damage are incorporated into our tendon model. Using a stochastic model of fiber repair, it is assumed that mechanically damaged fibers are repaired longer, whereas proteolytically damaged fibers are repaired shorter, relative to their pre-damage length. To study adaptation of tendon properties to applied load, our model musculotendon unit is a simplified three-component Hill-type model of the human Achilles-soleus unit. Our model results demonstrate that the geometric equilibrium state of the Achilles tendon can coincide with minimization of the total metabolic cost of muscle activation. The proposed tendon model independently predicts rates of collagen fiber turnover that are in general agreement with in vivo experimental measurements. While the computational model here only represents a first step in a new approach to understanding the complex process of tendon remodeling in vivo, given these findings, it appears likely that the proposed framework may itself provide a useful theoretical foundation for developing valuable qualitative and quantitative insights into tendon physiology and pathology.

Soleus Muscle as a Surrogate for Health Status in Human Heart Failure.

We propose the hypothesis that soleus muscle function may provide a surrogate measure of functional capacity in patients with heart failure. We summarize literature pertaining to skeletal muscle as a locus of fatigue and present our recent findings, using in vivo imaging in combination with biomechanical experimentation and modeling, to reveal novel structure-function relationships in chronic heart failure skeletal muscle and gait.

A comparison of haemolytic responses in fore-foot and rear-foot distance runners.

This study examined the haemolytic effects of an interval-based running task in fore-foot and rear-foot striking runners. Nineteen male distance runners (10 fore-foot, 9 rear-foot) completed 8 × 3 min repeats at 90% vVO2peak on a motorised treadmill. Pre- and post-exercise venous blood samples were analysed for serum haptoglobin to quantify the haemolytic response to running. Vertical ground reaction forces were also captured via a force plate beneath the treadmill belt. Haptoglobin levels were significantly decreased following exercise (P = 0.001) in both groups (but not between groups), suggesting that the running task created a haemolytic stress. The ground reaction force data showed strong effect sizes for a greater peak force (d = 1.20) and impulse (d = 1.37) in fore-foot runners, and a greater rate of force development (d = 2.74) in rear-foot runners. The lack of difference in haptoglobin response between groups may be explained by the trend for fore-foot runners to experience greater peak force and impulse during the stance phase of their running gait, potentially negating any impact of the greater rate of force development occurring from the rear-foot runners' heel strike. Neither type of runner (fore-foot or rear-foot) appears more susceptible to technique-related foot-strike haemolysis.

The Use of Geometric Morphometric Techniques to Identify Sexual Dimorphism in Gait.

This study attempts to apply geometric morphometric techniques for the analysis of 3D kinematic marker-based gait data. As a test, we attempted to identify sexual dimorphism during the stance phase of the gait cycle. Two techniques were used to try to identify differences in the way males and females walk without the results being affected by individual differences in body shape and size. Twenty-eight kinematic markers were placed on the torso and legs of 6 male and 8 female subjects, and the 3D time varying coordinates of the kinematic markers were recorded. The gait cycle trials were time-normalized to 61 frames representing the stance phase of gait, and the change in the shape of the configuration of kinematic markers was analyzed using principal components analysis to produce 'gait signatures' that characterize the kinematics of each individual. The variation in the gait signatures was analyzed with a further principal components analysis. These methods were able to detect significant sexual dimorphism even after the effects of sexual body shape and size differences were factored out. We discuss insights gained from performing this study which may be of value to others attempting to apply geometric morphometric methods to motion analysis.

Joint-level mechanics of the walk-to-run transition in humans.

Two commonly proposed mechanical explanations for the walk-to-run transition (WRT) include the prevention of muscular over-exertion (effort) and the minimization of peak musculoskeletal loads and thus injury risk. The purpose of this study was to address these hypotheses at a joint level by analysing the effect of speed on discrete lower-limb joint kinetic parameters in humans across a wide range of walking and running speeds including walking above and running below the WRT speed. Joint work, peak instantaneous joint power, and peak joint moments in the sagittal and frontal plane of the ankle, knee and hip from eight participants were collected for 10 walking speeds (30-120% of their WRT) and 10 running speeds (80-170% of their WRT) on a force plate instrumented treadmill. Of the parameters analysed, three satisfied our statistical criteria of the 'effort-load' hypothesis of the WRT. Mechanical parameters that provide an acute signal (peak moment and peak power) were more strongly associated with the gait transition than parameters that reflect the mechanical function across a portion of the stride. We found that both the ankle (peak instantaneous joint power during swing) and hip mechanics (peak instantaneous joint power and peak joint moments in stance) can influence the transition from walking to running in human locomotion and may represent a cascade of mechanical events beginning at the ankle and leading to an unfavourable compensation at the hip. Both the ankle and hip mechanisms may contribute to gait transition by lowering the muscular effort of running compared with walking at the WRT speed. Although few of the examined joint variables satisfied our hypothesis of the WRT, most showed a general marked increase when switching from walking to running across all speeds where both walking and running are possible, highlighting the fundamental differences in the mechanics of walking and running. While not eliciting the WRT per se, these variables may initiate the transition between stable walking and running patterns. Those variables that were invariant of gait were predominantly found in the swing phase.

Programmable mechanical stimulation influences tendon homeostasis in a bioreactor system.

Identification of functional programmable mechanical stimulation (PMS) on tendon not only provides the insight of the tendon homeostasis under physical/pathological condition, but also guides a better engineering strategy for tendon regeneration. The aims of the study are to design a bioreactor system with PMS to mimic the in vivo loading conditions, and to define the impact of different cyclic tensile strain on tendon. Rabbit Achilles tendons were loaded in the bioreactor with/without cyclic tensile loading (0.25 Hz for 8 h/day, 0-9% for 6 days). Tendons without loading lost its structure integrity as evidenced by disorientated collagen fiber, increased type III collagen expression, and increased cell apoptosis. Tendons with 3% of cyclic tensile loading had moderate matrix deterioration and elevated expression levels of MMP-1, 3, and 12, whilst exceeded loading regime of 9% caused massive rupture of collagen bundle. However, 6% of cyclic tensile strain was able to maintain the structural integrity and cellular function. Our data indicated that an optimal PMS is required to maintain the tendon homeostasis and there is only a narrow range of tensile strain that can induce the anabolic action. The clinical impact of this study is that optimized eccentric training program is needed to achieve maximum beneficial effects on chronic tendinopathy management.

Effects of plyometric training on achilles tendon properties and shuttle running during a simulated cricket batting innings.

The aim of this study was to determine whether intermittent shuttle running times (during a prolonged, simulated cricket batting innings) and Achilles tendon properties were affected by 8 weeks of plyometric training (PLYO, n = 7) or normal preseason (control [CON], n = 8). Turn (5-0-5-m agility) and 5-m sprint times were assessed using timing gates. Achilles tendon properties were determined using dynamometry, ultrasonography, and musculoskeletal geometry. Countermovement and squat jump heights were also assessed before and after training. Mean 5-0-5-m turn time did not significantly change in PLYO or CON (pre vs. post: 2.25 ± 0.08 vs. 2.22 ± 0.07 and 2.26 ± 0.06 vs. 2.25 ± 0.08 seconds, respectively). Mean 5-m sprint time did not significantly change in PLYO or CON (pre vs. post: 0.85 ± 0.02 vs. 0.84 ± 0.02 and 0.85 ± 0.03 vs. 0.85 ± 0.02 seconds, respectively). However, inferences from the smallest worthwhile change suggested that PLYO had a 51-72% chance of positive effects but only 6-15% chance of detrimental effects on shuttle running times. Jump heights only increased in PLYO (9.1-11.0%, p < 0.050). Achilles tendon mechanical properties (force, stiffness, elastic energy, strain, modulus) did not change in PLYO or CON. However, Achilles tendon cross-sectional area increased in PLYO (pre vs. post: 70 ± 7 vs. 79 ± 8 mm, p < 0.01) but not CON (77 ± 4 vs. 77 ± 5 mm, p > 0.050). In conclusion, plyometric training had possible benefits on intermittent shuttle running times and improved jump performance. Also, plyometric training increased tendon cross-sectional area, but further investigation is required to determine whether this translates to decreased injury risk.

Achilles tendon mechanical properties after both prolonged continuous running and prolonged intermittent shuttle running in cricket batting.

Effects of prolonged running on Achilles tendon properties were assessed after a 60 min treadmill run and 140 min intermittent shuttle running (simulated cricket batting innings). Before and after exercise, 11 participants performed ramp-up plantar flexions to maximum-voluntary-contraction before gradual relaxation. Muscle-tendon-junction displacement was measured with ultrasonography. Tendon force was estimated using dynamometry and a musculoskeletal model. Gradients of the ramp-up force-displacement curves fitted between 0-40% and 50-90% of the preexercise maximal force determined stiffness in the low- and high-force-range, respectively. Hysteresis was determined using the ramp-up and relaxation force-displacement curves and elastic energy storage from the area under the ramp-up curve. In simulated batting, correlations between tendon properties and shuttle times were also assessed. After both protocols, Achilles tendon force decreased (4% to 5%, P < .050), but there were no changes in stiffness, hysteresis, or elastic energy. In simulated batting, Achilles tendon force and stiffness were both correlated to mean turn and mean sprint times (r = -0.719 to -0.830, P < .050). Neither protocol resulted in fatigue-related changes in tendon properties, but higher tendon stiffness and plantar flexion force were related to faster turn and sprint times, possibly by improving force transmission and control of movement when decelerating and accelerating.

On the ascent: the soleus operating length is conserved to the ascending limb of the force-length curve across gait mechanics in humans.

The region over which skeletal muscles operate on their force-length (F-L) relationship is fundamental to the mechanics, control and economy of movement. Yet surprisingly little experimental data exist on normalized length operating ranges of muscle during human gait, or how they are modulated when mechanical demands (such as force output) change. Here we explored the soleus muscle (SOL) operating lengths experimentally in a group of healthy young adults by combining subject-specific F-L relationships with in vivo muscle imaging during gait. We tested whether modulation of operating lengths occurred between walking and running, two gaits that require different levels of force production and different muscle-tendon mechanics, and examined the relationship between optimal fascicle lengths (L(0)) and normalized operating lengths during these gaits. We found that the mean active muscle lengths reside predominantly on the ascending limbs of the F-L relationship in both gaits (walk, 0.70-0.94 L(0); run, 0.65-0.99 L(0)). Furthermore, the mean normalized muscle length at the time of the peak activation of the muscle was the same between the two gaits (0.88 L(0)). The active operating lengths were conserved, despite a fundamentally different fascicle strain pattern between walking (stretch-shorten cycle) and running (near continuous shortening). Taken together, these findings indicate that the SOL operating length is highly conserved, despite gait-dependent differences in muscle-tendon dynamics, and appear to be preferentially selected for stable force production compared with optimal force output (although length-dependent force capacity is high when maximal forces are expected to occur). Individuals with shorter L(0) undergo smaller absolute muscle excursions (P<0.05) so that the normalized length changes during walking and running remain independent of L(0). The correlation between L(0) and absolute length change was not explained on the basis of muscle moment arms or joint excursion, suggesting that regulation of muscle strain may occur via tendon stretch.

Running birds reveal secrets for legged robot design.

Recapitulating avian locomotion opens the door for simple and economical control of legged robots without sensory feedback systems.

Gait-specific energetics contributes to economical walking and running in emus and ostriches.

A widely held assumption is that metabolic rate (E(met)) during legged locomotion is linked to the mechanics of different gaits and this linkage helps explain the preferred speeds of animals in nature. However, despite several prominent exceptions, E(met) of walking and running vertebrates has been nearly uniformly characterized as increasing linearly with speed across all gaits. This description of locomotor energetics does not predict energetically optimal speeds for minimal cost of transport (E(cot)). We tested whether large bipedal ratite birds (emus and ostriches) have gait-specific energetics during walking and running similar to those found in humans. We found that during locomotion, emus showed a curvilinear relationship between E(met) and speed during walking, and both emus and ostriches demonstrated an abrupt change in the slope of E(met) versus speed at the gait transition with a linear increase during running. Similar to human locomotion, the minimum net E(cot) calculated after subtracting resting metabolism was lower in walking than in running in both species. However, the difference in net E(cot) between walking and running was less than is found in humans because of a greater change in the slope of E(met) versus speed at the gait transition, which lowers the cost of running for the avian bipeds. For emus, we also show that animals moving freely overground avoid a range of speeds surrounding the gait-transition speed within which the E(cot) is large. These data suggest that deviations from a linear relation of metabolic rate and speed and variations in transport costs with speed are more widespread than is often assumed, and provide new evidence that locomotor energetics influences the choice of speed in bipedal animals. The low cost of transport for walking is probably ecologically important for emus and ostriches because they spend the majority of their active day walking, and thus the energy used for locomotion is a large part of their daily energy budget.

Mechanisms producing coordinated function across the breadth of a large biarticular thigh muscle.

We examined the hypothesis that structural features of the iliotibialis lateralis pars postacetabularis (ILPO) in guinea fowl allow this large muscle to maintain equivalent function along its anterior-posterior axis. The ILPO, the largest muscle in the hindlimb of the guinea fowl, is a hip and knee extensor. The fascicles of the ILPO originate across a broad region of the ilium and ischium posterior to the hip. Its long posterior fascicles span the length of the thigh and insert directly on the patellar tendon complex. However, its anterior fascicles are shorter and insert on a narrow aponeurosis that forms a tendinous band along the anterior edge of the muscle and is connected distally to the patellar tendon. The biarticular ILPO is actively lengthened and then actively shortened during stance. The moment arm of the fascicles at the hip increases along the anterior to posterior axis, whereas the moment arm at the knee is constant for all fascicles. Using electromyography and sonomicrometry, we examined the activity and strain of posterior and anterior fascicles of the ILPO. The activation was not significantly different in the anterior and posterior fascicles. Although we found significant differences in active lengthening and shortening strain between the anterior and posterior fascicles, the differences were small. The majority of shortening strain is caused by hip extension and the inverse relationship between hip moment arm and fascicle length along the anterior-posterior axis was found to have a major role in ensuring similar shortening strain. However, because the knee moment arm is the same for all fascicles, knee flexion in early stance was predicted to produce much larger lengthening strains in the short anterior fascicles than our measured values at this location. We propose that active lengthening of the anterior fascicles was lower than predicted because the aponeurotic tendon of insertion of the anterior fascicles was stretched and only a portion of the lengthening had to be accommodated by the active muscle fascicles.

Understanding muscle energetics in locomotion: new modeling and experimental approaches.

Recent estimates of muscle energy consumption during locomotion, based on computational models and muscle blood flow measurements, demonstrate complex patterns of energy use across the gait cycle, which are further complicated when task demands change. A deeper understanding of muscle energetics in locomotion will benefit from efforts to more tightly integrate muscle-specific approaches with organismal measurements.

Performance in a simulated cricket batting innings (BATEX): reliability and discrimination between playing standards.

The reliability (test-retest) of running-between-the-wickets times and skill performance was assessed during a batting exercise (BATEX) simulation of 2 h 20 min duration that requires intermittent shuttle running. In addition, performance and physiological responses (heart rate, sweat rate, rating of perceived exertion, blood lactate concentration) were compared between high- and low-grade district club batsmen (n = 22, mean ± s: age 20 ± 2 years, mass 73.4 ± 8.5 kg). Running-between-the-wickets performance was assessed with an infra-red timing system (Swift, Australia) by sampling a 5-m time for the middle section of the straight-line sprints (singles) and the time to complete 5 m in and out of the turn (5-0-5-m turn time). Skill performance was rated as a percentage for good bat-ball contacts. Coefficients of variation for running-between-the-wickets performance and percentage of good bat-ball contacts were both <5%. Percentage of good bat-ball contacts was greater in the high- than low-grade batsmen (70 ± 8 vs. 58 ± 9%, P = 0.01). All other variables were similar between grades. Running-between-the-wickets and skill-performance measures during the BATEX simulation were reliable, thus it can be used in future research.

Movement patterns and physical strain during a novel, simulated cricket batting innings (BATEX).

A simulated cricket batting innings was developed to replicate the physical demands of scoring a century during One-Day International cricket. The simulated innings requires running-between-the-wickets across six 5-over stages, each of 21 min duration. To validate whether the simulated batting innings is reflective of One-Day International batting, movement patterns were collected using a global positioning system (GPS) and compared with previous research. In addition, indicators of physical strain were recorded (heart rate, jump heights, sweat loss, tympanic temperature). Nine club cricketers (mean ± s: age 20 ± 3 years; body mass 79.5 ± 7.9 kg) performed the simulated innings outdoors. There was a moderate trend for distance covered in the simulated innings to be less than that during One-Day batting (2171 ± 157 vs. 2476 ± 631 m · h⁻¹; effect size = 0.78). This difference was largely explained by a strong trend for less distance covered walking in the simulated innings than in One-Day batting (1359 ± 157 vs. 1604 ± 438 m · h⁻¹; effect size = 1.61). However, there was a marked trend for distance covered both striding and sprinting to be greater in the simulated innings than in One-Day batting (effect size > 1.2). Practically, the simulated batting innings may be used for match-realistic physical training and as a research protocol to assess the demands of prolonged, high-intensity cricket batting.

Influence of stretching and warm-up on Achilles tendon material properties.

BACKGROUND: Controversy exists on stretching and warm-up in injury prevention. We hypothesized that warm up has a greater effect on Achilles tendon biomechanics than static stretching. This study investigated static stretching and warm-up on Achilles tendon biomechanics in recreational athletes, in vivo. MATERIALS AND METHODS: Ten active, healthy subjects, 5 males, 5 females, With a mean age of 22.9 years with no previous Achilles tendon injuries were recruited. Typical stretching and warm-up routines were created. Testing was performed in a randomized cross-over design. A custom-built dynamometer was utilized to perform controlled isometric plantarflexion. A low profile ultrasound probe was utilized to visualize the musculotendinous junction of the medial gastrocnemius. An eight-camera motion capture system was used to capture ankle motion. Custom software calculated Achilles tendon biomechanics. RESULTS: Achilles tendon force production was consistent. No statistically significant differences were detected in stretch, stiffness, and strain between pre-, post-stretching, and post-warm-up interventions. CONCLUSION: Stretching or warm-up alone, and combined did not demonstrate statistically significant differences. Stretching and warm-up may have an equivalent effect on Achilles tendon biomechanics. Prolonged and more intense protocols may be required for changes to occur. CLINICAL RELEVANCE: Stretching and warm-up of the Achilles before exercise are commonly practiced. Investigating the effect of stretching and warm-up may shed light on potential injury prevention.