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

Tendon mechanical properties respond to altered load in adults, but how load history during growth affects adult tendon properties remains unclear. To address this question, we adopted an avian model in which we altered the mechanical load environment across the growth span. Animals were divided at 2 weeks of age into three groups: (1) an exercise control group given the opportunity to perform high-acceleration movements (EXE, n = 8); (2) a sedentary group restricted from high-intensity exercise (RES, n = 8); and (3) a sedentary group also restricted from high-intensity exercise and in which the gastrocnemius muscles were partially paralyzed using repeated bouts of botulinum toxin-A injections (RES-BTX, n = 8). Video analysis of bird movement confirmed the restrictions eliminated high-intensity exercise and did not alter time spent walking and sitting between groups. At skeletal maturity (33–35 weeks) animals were sacrificed for analysis, consisting of high-field MRI and material load testing, of both the entire free Achilles tendon and the tendon at the bone-tendon junction. Free tendon stiffness, modulus, and hysteresis were unaffected by variation in load environment. Further, the bone-tendon junction cross-sectional area, stress, and strain were also unaffected by variations in load environment. These results suggest that: (a) a baseline level of low-intensity activity (standing and walking) may be sufficient to maintain tendon growth; and (b) if this lower threshold of tendon load is met, non-mechanical mediated tendon growth may override the load-induced mechanotransduction signal attributed to tendon remodeling in adults of the same species. These results are important for understanding of musculoskeletal function and tendon health in growing individuals.

Sprinters have been found to possess longer muscle fascicles than non-sprinters, which is thought to be beneficial for high-acceleration movements based on muscle force-length-velocity properties. However, it is unknown if their morphology is a result of genetics or training during growth. To explore the influence of training during growth, thirty guinea fowl (Numida meleagris) were split into exercise and sedentary groups. Exercise birds were housed in a large pen and underwent high-acceleration training during their growth period (age 4–14 weeks), while sedentary birds were housed in small pens to restrict movement. Morphological analyses (muscle mass, PCSA, optimal fascicle length, pennation angle) of a hip extensor muscle (ILPO) and plantarflexor muscle (LG), which differ in architecture and function during running, were performed post-mortem. Muscle mass for both ILPO and LG was not different between the two groups. Exercise birds were found to have ∼12% and ∼14% longer optimal fascicle lengths in ILPO and LG, respectively, than the sedentary group despite having ∼3% shorter limbs. From this study we can conclude that optimal fascicle lengths can increase as a result of high-acceleration training during growth. This increase in optimal fascicle length appears to occur irrespective of muscle architecture and in the absence of a change in muscle mass. Our findings suggest high-acceleration training during growth results in muscles that prioritize adaptations for lower strain and shortening velocity over isometric strength. Thus, the adaptations observed suggest these muscles produce higher force during dynamic contractions, which is beneficial for movements requiring large power outputs.

Similar to most biological tissues, the biomechanical, and functional characteristics of the Achilles tendon are closely related to its composition and microstructure. It is commonly reported that type I collagen is the predominant component of tendons and is mainly responsible for the tissue's function. Although elastin has been found in varying proportions in other connective tissues, previous studies report that tendons contain very small quantities of elastin. However, the morphology and the microstructural relationship among the elastic fibres, collagen, and cells in tendon tissue have not been well examined. We hypothesize the elastic fibres, as another fibrillar component in the extracellular matrix, have a unique role in mechanical function and microstructural arrangement in Achilles tendons. It has been shown that elastic fibres present a close connection with the tenocytes. The close relationship of the three components has been revealed as a distinct, integrated and complex microstructural network. Notably, a "spiral" structure within fibril bundles in Achilles tendons was observed in some samples in specialized regions. This study substantiates the hierarchical system of the spatial microstructure of tendon, including the mapping of collagen, elastin and tenocytes, with 3-dimensional confocal images. © 2017 Orthopaedic Research Society. Published by Wiley Periodicals, Inc. J Orthop Res 35:1203-1214, 2017.

Decreases in activity levels in children worldwide are feared to have long-term health repercussions. Yet, because of the difficulty of performing controlled long-term studies in humans, we do not yet understand how decreases in childhood activity influence adult functional capacity. Here, in an avian bipedal model, we evaluated the elimination of all high-intensity activity during growth on adult performance. We evaluated three alternative hypotheses: Elimination of high-intensity activity 1) does not influence adult function, 2) results in task-specific deficits in adulthood, or 3) results in deficits that generalize across locomotor tasks. We found that animals restricted from jumping and sprinting during growth showed detriments as adults in maximal jump performance in comparison to controls, but did not require more metabolic energy during steady-state running or standing. From this, we conclude that functional deficits from elimination of high-intensity exercise are task specific and do not generalize across all locomotor functions.NEW & NOTEWORTHY Decreasing childhood activity levels are feared to have long-term health repercussions, but testing this hypothesis is hampered by restrictions of human experimentation. Here, in a bipedal animal model, we examine how the elimination of high-intensity activity during all of maturation influences adult locomotor capacity. We found restricted activity during growth reduced mechanical power capacity but not submaximal metabolic cost. This suggests that reduced childhood activity may result in task-specific, rather than generalized locomotor deficits.

Minimization of metabolic energy is considered a fundamental principle of human locomotion, as demonstrated by an alignment between the preferred walking speed (PWS) and the speed incurring the lowest metabolic cost of transport. We aimed to (i) simultaneously disrupt metabolic cost and an alternate acute task requirement, namely speed error regulation, and (ii) assess whether the PWS could be explained on the basis of either optimality criterion in this new performance and energetic landscape. Healthy adults ( N = 21) walked on an instrumented treadmill under normal conditions and, while negotiating a continuous gait perturbation, imposed leg-length asymmetry. Oxygen consumption, motion capture data and ground reaction forces were continuously recorded for each condition at speeds ranging from 0.6 to 1.8 m s −1 , including the PWS. Both metabolic and speed regulation measures were disrupted by the perturbation ( p < 0.05). Perturbed PWS selection did not exhibit energetic prioritization (although we find some indication of energy minimization after motor adaptation). Similarly, PWS selection did not support prioritization of speed error regulation, which was found to be independent of speed in both conditions. It appears that, during acute exposure to a mechanical gait perturbation of imposed leg-length asymmetry, humans minimize neither energetic cost nor speed regulation errors. Despite the abundance of evidence pointing to energy minimization during normal, steady-state gait, this may not extend acutely to perturbed gait. Understanding how the nervous system acutely controls gait perturbations requires further research that embraces multi-objective control paradigms.

A muscle's performance is influenced by where it operates on its force-length (F-L) curve. Here we explore how activation and tendon compliance interact to influence muscle operating lengths and force-generating capacity. To study this, we built a musculoskeletal model of the lower limb of the guinea fowl and simulated the F-L operating range during fixed-end fixed-posture contractions for 39 actuators under thousands of combinations of activation and posture using three different muscle models: Muscles with non-compliant tendons, muscles with compliant tendons but no activation-dependent shift in optimal fiber length (L0), and muscles with both compliant tendons and activation-dependent shifts in L0. We found that activation-dependent effects altered muscle fiber lengths up to 40% and increased or decreased force capacity by up to 50% during fixed-end contractions. Typically, activation-compliance effects reduce muscle force and are dominated by the effects of tendon compliance at high activations. At low activation, however, activation-dependent shifts in L0 are equally important and can result in relative force changes for low compliance muscles of up to 60%. There are regions of the F-L curve in which muscles are most sensitive to compliance and there are troughs of influence where these factors have little effect. These regions are hard to predict, though, because the magnitude and location of these areas of high and low sensitivity shift with compliance level. In this study we provide a map for when these effects will meaningfully influence force capacity and an example of their contributions to force production during a static task, namely standing.A Interação de Conformidade e Ativação na Faixa de Operação Força-Comprimento e Capacidade de Geração de Força do Músculo Esquelético: Um Estudo Computacional Usando um Modelo Musculoesquelético de Galinhas-D’angola O desempenho muscular é influenciado por onde ele opera na sua curva de força-comprimento. Aqui, exploramos como a ativação e a conformidade do tendão interagem para influenciar os comprimentos musculares e a capacidade de geração de força. Para estudar isso, construímos um modelo musculoesquelético do membro inferior da galinha-d’angola e simulamos a faixa de operação força-comprimento durante contrações fixas de postura e extremidade para 39 atuadores sob milhares de combinações de ativação e postura usando três modelos musculares diferentes: músculos com tendões não-complacentes, músculos com tendões complacentes, mas sem desvio dependente de ativação no comprimento ideal de fibra (L0), e músculos com tendões complacentes e desvios dependentes de ativação em L0. Descobrimos que os efeitos dependentes da ativação alteraram os comprimentos da fibra muscular em até 40% e aumentaram ou diminuíram a capacidade de força em até 50% durante as contrações de extremidade fixas. Normalmente, os efeitos de ativação e conformidade reduzem a força muscular e são dominados pelos efeitos de complacência do tendão em altas ativações. Em baixa ativação, no entanto, desvios dependentes de ativação em L0 são igualmente importantes e podem resultar em mudanças de força relativas de até 60% para músculos de baixa complacência. Existem regiões da curva de força-comprimento em que os músculos são mais sensíveis à complacência e há baixas de influência onde esses fatores têm pouco efeito. Essas regiões são difíceis de prever porque a magnitude e a localização dessas áreas de alta e baixa sensibilidade mudam com o nível de conformidade. Neste estudo, fornecemos um mapa para quando esses efeitos influenciarão significativamente a capacidade de força e um exemplo de suas contribuições para a produção de forças durante uma tarefa estática, ou seja, em pé. Translated to Portuguese by G. Sobral (gabisobral@gmail.com).

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

Wearable robotics offers a unique opportunity to explore how biological systems interface with engineered parts. But, due to a gap in understanding of the underlying biological mechanisms at work, the state of the art in design and development is a sophisticated form of automated trial and error. Progress is hampered by the difficulty of assessing the direct impact of wearable robots on underlying muscles, tendons and bones during human experimentation. While animal models have provided an experimental platform to explore other biological mechanisms, as of yet, no animal model of a wearable robot during locomotion has been developed. To fill this gap, we have built the first ever wearable robotic device for a freely-Iocomoting, non-human, bipedal animal (Numida melaegris = Guinea fowl), a species whose gait closely mirrors human locomotion mechanics. We found that a spring-loaded soft-exosuit that passively augments the energy stored in distal tendons was both well tolerated and provided consistent torques. Preliminary data showed birds systematically change their kinematics in response to changes to exo-suit spring stiffness, adjusting the timing but not magnitude of the assistive torques. This animal model for wearable robotics allows experiments up and down the broader spatiotemporal scale that are not currently possible in humans. With it we can address questions from short-term adaptations in musculoskeletal dynamics within a single step to broader behavioral and physical changes that come with long term use.

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 (WLK) to a variety of running velocities (RUN; ranging from a sprint to a velocity slower than WLK). WRTs to fast RUNs 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 of WLK 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 RUN velocity, 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 RUNs, 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.

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.

Physiotherapy is one of the effective treatments for tendinopathy, whereby symptoms are relieved by changing the biomechanical environment of the pathological tendon. However, the underlying mechanism remains unclear. In this study, we first established a model of progressive tendinopathy-like degeneration in the rabbit Achilles. Following ex vivo loading deprivation culture in a bioreactor system for 6 and 12 days, tendons exhibited progressive degenerative changes, abnormal collagen type III production, increased cell apoptosis, and weakened mechanical properties. When intervention was applied at day 7 for another 6 days by using cyclic tensile mechanical stimulation (6% strain, 0.25 Hz, 8 h/day) in a bioreactor, the pathological changes and mechanical properties were almost restored to levels seen in healthy tendon. Our results indicated that a proper biomechanical environment was able to rescue early-stage pathological changes by increased collagen type I production, decreased collagen degradation and cell apoptosis. The ex vivo model developed in this study allows systematic study on the effect of mechanical stimulation on tendon biology.

Reduced walking capacity, a hallmark of chronic heart failure (CHF), is strongly correlated with hospitalization and morbidity. The aim of this work was to perform a detailed biomechanical gait analysis to better identify mechanisms underlying reduced walking capacity in CHF. Inverse dynamic analyses were conducted in CHF patients and age- and exercise level-matched control subjects on an instrumented treadmill at self-selected treadmill walking speeds and at speeds representing +20% and -20% of the subjects' preferred speed. Surprisingly, no difference in preferred speed was observed between groups, possibly explained by an optimization of the mechanical cost of transport in both groups (the mechanical cost to travel a given distance; J/kg/m). The majority of limb kinematics and kinetics were also similar between groups, with the exception of greater ankle dorsiflexion angles during stance in CHF. Nevertheless, over two times greater ankle plantarflexion work during stance and per distance traveled is required for a given triceps surae muscle volume in CHF patients. This, together with a greater reliance on the ankle compared to the hip to power walking in CHF patients, especially at faster speeds, may contribute to the earlier onset of fatigue in CHF patients. This observation also helps explain the high correlation between triceps surae muscle volume and exercise capacity that has previously been reported in CHF. Considering the key role played by the plantarflexors in powering walking and their association with exercise capacity, our findings strongly suggest that exercise-based rehabilitation in CHF should not omit the ankle muscle group.

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.

Computational modeling of tendon lags the development of computational models for other tissues. A major bottleneck in the development of realistic computational models for Achilles tendon is the absence of detailed conceptual and theoretical models as to how the tissue actually functions. Without the conceptual models to provide a theoretical framework to guide the development and integration of multiscale computational models, modeling of the Achilles tendon to date has tended to be piecemeal and focused on specific mechanical or biochemical issues. In this paper, we present a new conceptual model of Achilles tendon tissue homeostasis, and discuss this model in terms of existing computational models of tendon. This approach has the benefits of structuring the research on relevant computational modeling to date, while allowing us to identify new computational models requiring development. The critically important functional issue for tendon is that it is continually damaged during use and so has to be repaired. From this follows the centrally important issue of homeostasis of the load carrying collagen fibrils within the collagen fibers of the Achilles tendon. Collagen fibrils may be damaged mechanically-by loading, or damaged biochemically-by proteases. Upon reviewing existing computational models within this conceptual framework of the Achilles tendon structure and function, we demonstrate that a great deal of theoretical and experimental research remains to be done before there are reliably predictive multiscale computational model of Achilles tendon in health and disease.

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%, p0.050). 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, p0.01) but not CON (77 ± 4 vs. 77 ± 5 mm, p0.050). 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.

Tendon and ligament injury is a worldwide health problem, but the treatment options remain limited. Tendon and ligament engineering might provide an alternative tissue source for the surgical replacement of injured tendon. A bioreactor provides a controllable environment enabling the systematic study of specific biological, biochemical, and biomechanical requirements to design and manufacture engineered tendon/ligament tissue. Furthermore, the tendon/ligament bioreactor system can provide a suitable culture environment, which mimics the dynamics of the in vivo environment for tendon/ligament maturation. For clinical settings, bioreactors also have the advantages of less-contamination risk, high reproducibility of cell propagation by minimizing manual operation, and a consistent end product. In this review, we identify the key components, design preferences, and criteria that are required for the development of an ideal bioreactor for engineering tendons and ligaments.

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.

This study combines experimental-based and model-based methodologies for accessing in vivo musculoskeletal function in healthy individuals. We use ultrasound and dynamometer technologies to derive subject-specific muscle parameters including muscle isometric force, optimal fiber length and tendon slack length. We then assess the impact of subject-specificity on the electromyography-driven simulation of walking of the composite musculoskeletal system.

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.

Owing to their cursorial background, ostriches (Struthio camelus) walk and run with high metabolic economy, can reach very fast running speeds and quickly execute cutting manoeuvres. These capabilities are believed to be a result of their ability to coordinate muscles to take advantage of specialized passive limb structures. This study aimed to infer the functional roles of ostrich pelvic limb muscles during gait. Existing gait data were combined with a newly developed musculoskeletal model to generate simulations of ostrich walking and running that predict muscle excitations, force and mechanical work. Consistent with previous avian electromyography studies, predicted excitation patterns showed that individual muscles tended to be excited primarily during only stance or swing. Work and force estimates show that ostrich gaits are partially hip-driven with the bi-articular hip–knee muscles driving stance mechanics. Conversely, the knee extensors acted as brakes, absorbing energy. The digital extensors generated large amounts of both negative and positive mechanical work, with increased magnitudes during running, providing further evidence that ostriches make extensive use of tendinous elastic energy storage to improve economy. The simulations also highlight the need to carefully consider non-muscular soft tissues that may play a role in ostrich gait.

SUMMARY 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.

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 both5%. 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.

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.

A widely held assumption is that metabolic rate (Ėmet) 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,Ėmetof 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 (Ecot). 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 betweenĖmetand speed during walking, and both emus and ostriches demonstrated an abrupt change in the slope ofĖmetversus speed at the gait transition with a linear increase during running. Similar to human locomotion, the minimum netEcotcalculated after subtracting resting metabolism was lower in walking than in running in both species. However, the difference in netEcotbetween walking and running was less than is found in humans because of a greater change in the slope ofĖmetversus 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 theEcotis 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.

Understanding the mechanical determinants of the energy cost of limb swing is crucial for refining our models of locomotor energetics, as well as improving treatments for those suffering from impaired limb-swing mechanics. In this study, we use guinea fowl ( Numida meleagris) as a model to explore whether mechanical work at the joints explains limb-swing energy use by combining inverse dynamic modeling and muscle-specific energetics from blood flow measurements. We found that the overall efficiencies of the limb swing increased markedly from walking (3%) to fast running (17%) and are well below the usually accepted maximum efficiency of muscle, except at the fastest speeds recorded. The estimated efficiency of a single muscle used during ankle flexion (tibialis cranialis) parallels that of the total limb-swing efficiency (3% walking, 15% fast running). Taken together, these findings do not support the hypothesis that joint work is the major determinant of limb-swing energy use across the animal's speed range and warn against making simple predictions of energy use based on joint mechanical work. To understand limb-swing energy use, mechanical functions other than accelerating the limb segments need to be explored, including isometric force production and muscle work arising from active and passive antagonist muscle forces.

SUMMARY Although locomotor kinematics in walking and running birds have been examined in studies exploring many biological aspects of bipedalism, these studies have been largely limited to two-dimensional analyses. Incorporating a five-segment, 17 degree-of-freedom (d.f.) kinematic model of the ostrich hind limb developed from anatomical specimens, we quantified the three-dimensional(3-D) joint axis alignment and joint kinematics during running (at ∼3.3 m s–1) in the largest avian biped, the ostrich. Our analysis revealed that the majority of the segment motion during running in the ostrich occurs in flexion/extension. Importantly, however, the alignment of the average flexion/extension helical axes of the knee and ankle are rotated externally to the direction of travel (37° and 21°, respectively) so that pure flexion and extension at the knee will act to adduct and adbuct the tibiotarsus relative to the plane of movement, and pure flexion and extension at the ankle will act to abduct and adduct the tarsometatarsus relative to the plane of movement. This feature of the limb anatomy appears to provide the major lateral (non-sagittal) displacement of the lower limb necessary for steering the swinging limb clear of the stance limb and replaces what would otherwise require greater adduction/abduction and/or internal/external rotation, allowing for less complex joints, musculoskeletal geometry and neuromuscular control. Significant rotation about the joints'non-flexion/extension axes nevertheless occurs over the running stride. In particular, hip abduction and knee internal/external and varus/valgus motion may further facilitate limb clearance during the swing phase, and substantial non-flexion/extension movement at the knee is also observed during stance. Measurement of 3-D segment and joint motion in birds will be aided by the use of functionally determined axes of rotation rather than assumed axes, proving important when interpreting the biomechanics and motor control of avian bipedalism.

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.

Physiotherapy is one of the effective treatments for tendinopathy, whereby symptoms are relieved by changing the biomechanical environment of the pathological tendon. However, the underlying mechanism remains unclear. In this study, we first established a model of progressive tendinopathy-like degeneration in the rabbit Achilles. Following ex vivo loading deprivation culture in a bioreactor system for 6 and 12 days, tendons exhibited progressive degenerative changes, abnormal collagen type III production, increased cell apoptosis, and weakened mechanical properties. When intervention was applied at day 7 for another 6 days by using cyclic tensile mechanical stimulation (6% strain, 0.25 Hz, 8 h/day) in a bioreactor, the pathological changes and mechanical properties were almost restored to levels seen in healthy tendon. Our results indicated that a proper biomechanical environment was able to rescue early-stage pathological changes by increased collagen type I production, decreased collagen degradation and cell apoptosis. The ex vivo model developed in this study allows systematic study on the effect of mechanical stimulation on tendon biology.

SUMMARYWe examined the energetic cost of loading the trunk or distal portion of the leg in walking and running guinea fowl (Numida meleagris). These different loading regimes were designed to separately influence the energy use by muscles used during the stance and swing phases of the stride. Metabolic rate, estimated from oxygen consumption, was measured while birds locomoted on a motorized treadmill at speeds from 0.5 to 2.0 m s-1, either unloaded, or with a mass equivalent to 23% of their body mass carried on their backs, or with masses equal to approximately 2.5% of their body mass attached to each tarsometatarsal segment. In separate experiments, we also measured the duration of stance and swing in unloaded, trunk-loaded, or limb-loaded birds. In the unloaded and limb-loaded birds, we also calculated the mechanical energy of the tarsometatarsal segment throughout the stride.Trunk and limb loads caused similar increases in metabolic rate. During trunk loading, the net metabolic rate (gross metabolic rate - resting metabolic rate) increased by 17% above the unloaded value across all speeds. This percentage increase is less than has been found in most studies of humans and other mammals. The economical load carriage of guinea fowl is consistent with predictions based on the relative cost of the stance and swing phases of the stride in this species. However, the available comparative data and considerations of the factors that determine the cost of carrying extra mass lead us to the conclusion that the cost of load carrying is unlikely to be a reliable indicator of the distribution of energy use in stance and swing. Both loading regimes caused small changes in the swing and/or stance durations, but these changes were less than 10%.Loading the tarsometatarsal segment increased its segmental energy by 4.1 times and the segmental mechanical power averaged over the stride by 3.8 times. The increases in metabolism associated with limb loading appear to be linked to the increases in mechanical power. The delta efficiency (change in mechanical power divided by the change in metabolic power) of producing this power increased from 11% in walking to approximately 25% in running. Although tarsometatarsal loading was designed to increase the mechanical energy during swing phase, 40% of the increase in segmental energy occurred during late stance. Thus, the increased energy demand of distal limb loading in guinea fowl is predicted to cause increases in energy use by both stance- and swing-phase muscles.

It has been argued that minimization of metabolic-energy costs is a primary determinant of gait selection in terrestrial animals. This view is based predominantly on data from humans and horses, which have been shown to choose the most economical gait (walking, running, galloping) for any given speed. It is not certain whether a minimization of metabolic costs is associated with the selection of other prevalent forms of terrestrial gaits, such as grounded running (a widespread gait in birds). Using biomechanical and metabolic measurements of four ostriches moving on a treadmill over a range of speeds from 0.8 to 6.7 m s(-1), we reveal here that the selection of walking or grounded running at intermediate speeds also favours a reduction in the metabolic cost of locomotion. This gait transition is characterized by a shift in locomotor kinetics from an inverted-pendulum gait to a bouncing gait that lacks an aerial phase. By contrast, when the ostrich adopts an aerial-running gait at faster speeds, there are no abrupt transitions in mechanical parameters or in the metabolic cost of locomotion. These data suggest a continuum between grounded and aerial running, indicating that they belong to the same locomotor paradigm.

Two commonly proposed mechanical explanations for the WRT include the prevention of muscular over-exertion (effort) and the minimisation 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 8 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 to 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.

AB Purpose: To better understand the mechanical factors differentiating forefoot and rearfoot strike (RFS) running, as well as the mechanical consequences of switching techniques, we assessed lower limb joint kinetics in habitual and imposed techniques in both groups. Methods: All participants performed both RFS and forefoot strike (FFS) techniques on an instrumented treadmill at 4.5 m[middle dot]s-1 while force and kinematic data were collected. Results: Total (sum of ankle, knee, and hip) lower limb work and average power did not differ between habitual RFS and FFS runners. However, moments, negative work and negative instantaneous and average power during stance were greater at the knee in RFS and at the ankle in FFS techniques. When habitual RFS runners switched to an imposed FFS, they were able to replicate the sagittal plane mechanics of a habitual FFS; however, the ankle internal rotation moment was increased by 33%, whereas the knee abduction moments were not reduced, remaining 48.5% higher than a habitual FFS. In addition, total positive and negative lower limb average power was increased by 17% and 9%, respectively. When habitual FFS runners switched to an imposed RFS, they were able to match the mechanics of habitual RFS runners with the exception of knee abduction moments, which remained 38% lower than a habitual RFS and, surprisingly, a reduction of total lower limb positive average power of 10.5%. Conclusions: There appears to be no clear overall mechanical advantage of a habitual FFS or RFS. Switching techniques may have different injury implications given the altered distribution in loading between joints but should be weighed against the overall effects on limb mechanics; adopting an imposed RFS may prove the most beneficial given the absence of any clear mechanical performance decrements

The biomechanical roles of skeletal muscle and their tendons are diverse. Perhaps most intuitively, muscle is regarded as a biological ‘motor’ that provides the work required for accelerating the body and overcoming aero- and hydrodynamic forces. With detailed biomechanical analyses, more intricate roles of the muscle-tendon unit have been uncovered, ranging from energy recyclers, to shock absorbers and capacitors. The functional scope of muscle-tendon tissue makes it an attractive choice for exploring bio-machine integration. Research and cross-disciplinary collaboration at SymbioticA offers a testbed for scientific and artistic exploration into engineered muscle-tendon constructs and the broader philosophical debate surrounding their place in ‘semi-living’ machine systems.

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., Author Summary It is now widely acknowledged that tendon plays a vital role in locomotion, while experiments have revealed that tendon is much more metabolically active than previously believed. There have been increasing numbers of papers describing the responses of tenocytes to mechanical loading and speculation about the origins of tendinopathy, but to date there is currently no basic theoretical framework describing how tendon maintains tissue homeostasis consistent with the experimental data, or indeed how tendon adapts to its environmental load conditions. Based on established biological principles of tendon damage and repair, for the first time we develop a dynamic model of tendon homeostasis that is capable of adaptation. We show that for a model soleus musculotendon unit with muscle fiber length kept constant, our model tendon is ‘capable’ of dynamically adjusting itself to find a stable equilibrium tendon geometry, which coincides with minimum metabolic cost of muscle activation. This new theoretical framework for tendon homeostasis and adaptation offers the possibility of refocusing research in basic and clinical science.

Elastic energy returned from passive-elastic structures of the lower limb is fundamental in lowering the mechanical demand on muscles during running. The purpose of this study was to investigate the two length-modulating mechanisms of the plantar fascia, namely medial longitudinal arch compression and metatarsophalangeal joint (MPJ) excursion, and to determine how these mechanisms modulate strain, and thus elastic energy storage/return of the plantar fascia during running. Eighteen runners (9 forefoot and 9 rearfoot strike) performed three treadmill running trials; unrestricted shod, shod with restricted arch compression (via an orthotic-style insert), and barefoot. Three-dimensional motion capture and ground reaction force data were used to calculate lower limb kinematics and kinetics including MPJ angles, moments, powers and work. Estimates of plantar fascia strain due to arch compression and MPJ excursion were derived using a geometric model of the arch and a subject-specific musculoskeletal model of the plantar fascia, respectively. The plantar fascia exhibited a typical elastic stretch-shortening cycle with the majority of strain generated via arch compression. This strategy was similar in fore- and rear-foot strike runners. Restricting arch compression, and hence the elastic-spring function of the arch, was not compensated for by an increase in MPJ-derived strain. In the second half of stance the plantar fascia was found to transfer energy between the MPJ (energy absorption) and the arch (energy production during recoil). This previously unreported energy transfer mechanism reduces the strain required by the plantar fascia in generating useful positive mechanical work at the arch during running.

Summary 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 explore 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 test whether modulation of operating lengths occurs between walking and running, two gaits that require different levels of force production and different muscle-tendon mechanics, and examine the relationship between optimal fascicle lengths (L0) 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 (0.70 - 0.94 L0, walk; 0.65 - 0.99 L0, run). Furthermore, the mean normalized muscle length at the time of the muscle's peak activation was the same between the two gaits (0.88 L0). 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-dependant differences in muscle-tendon dynamics, and appear to be preferentially selected for stable force production compared to optimal force output (although length-dependent force capacity is high when maximal forces are expected to occur). Individuals with shorter L0 undergo smaller absolute muscle excursions (p

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 size1.2). 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.

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.

SUMMARYThe alleged high net energy cost of running and low net energy cost of walking in humans have played an important role in the interpretation of the evolution of human bipedalism and the biomechanical determinants of the metabolic cost of locomotion. This study re-explores how the net metabolic energy cost of running and walking (J kg–1m–1) in humans compares to that of animals of similar mass using new allometric analyses of previously published data. Firstly, this study shows that the use of the slope of the regression between the rate of energy expenditure and speed to calculate the net energy cost of locomotion overestimates the net cost of human running. Also, the net energy cost of human running is only 17% higher than that predicted based on their mass. This value is not exceptional given that over a quarter of the previously examined mammals and birds have a net energy cost of running that is 17% or more above their allometrically predicted value. Using a new allometric equation for the net energy cost of walking, this study also shows that human walking is 20%less expensive than predicted for their mass. Of the animals used to generate this equation, 25% have a relatively lower net cost of walking compared with their allometrically predicted value. This new walking allometric analysis also indicates that the scaling of the net energy cost of locomotion with body mass is gait dependent. In conclusion, the net costs of running and walking in humans are moderately different from those predicted from allometry and are not remarkable for an animal of its size.

Active video games that require physical exertion during game play have been shown to confer health benefits. Typically, energy expended during game play is measured using devices attached to players, such as accelerometers, or portable gas analyzers. Since 2010, active video gaming technology incorporates marker-less motion capture devices to simulate human movement into game play. Using the Kinect Sensor and Microsoft SDK this research aimed to estimate the mechanical work performed by the human body and estimate subsequent metabolic energy using predictive algorithmic models. Nineteen University students participated in a repeated measures experiment performing four fundamental movements (arm swings, standing jumps, body-weight squats, and jumping jacks). Metabolic energy was captured using a Cortex Metamax 3B automated gas analysis system with mechanical movement captured by the combined motion data from two Kinect cameras. Estimations of the body segment properties, such as segment mass, length, centre of mass position, and radius of gyration, were calculated from the Zatsiorsky-Seluyanov's equations of de Leva, with adjustment made for posture cost. GPML toolbox implementation of the Gaussian Process Regression, a locally weighted k-Nearest Neighbour Regression, and a linear regression technique were evaluated for their performance on predicting the metabolic cost from new feature vectors. The experimental results show that Gaussian Process Regression outperformed the other two techniques by a small margin. This study demonstrated that physical activity energy expenditure during exercise, using the Kinect camera as a motion capture system, can be estimated from segmental mechanical work. Estimates for high-energy activities, such as standing jumps and jumping jacks, can be made accurately, but for low-energy activities, such as squatting, the posture of static poses should be considered as a contributing factor. When translated into the active video gaming environment, the results could be incorporated into game play to more accurately control the energy expenditure requirements.

SUMMARYUphill running requires more energy than level running at the same speed,largely due to the additional mechanical work of elevating the body weight. We explored the distribution of energy use among the leg muscles of guinea fowl running on the level and uphill using both organismal energy expenditure(oxygen consumption) and muscle blood flow measurements. We tested each bird under four conditions: (1) rest, (2) a moderate-speed level run at 1.5 m s–1, (3) an incline run at 1.5 m s–1 with a 15% gradient and (4) a fast level run at a speed eliciting the same metabolic rate as did running at a 15% gradient at 1.5 m s–1(2.28–2.39 m s–1). The organismal energy expenditure increased by 30% between the moderate-speed level run and both the fast level run and the incline run, and was matched by a proportional increase in total blood flow to the leg muscles. We found that blood flow increased significantly to nearly all the leg muscles between the moderate-speed level run and the incline run. However, the increase in flow was distributed unevenly across the leg muscles, with just three muscles being responsible for over 50% of the total increase in blood flow during uphill running. Three muscles showed significant increases in blood flow with increased incline but not with an increase in speed. Increasing the volume of active muscle may explain why in a previous study a higher maximal rate of oxygen consumption was measured during uphill running. The majority of the increase in energy expenditure between level and incline running was used in stance-phase muscles. Proximal stance-phase extensor muscles with parallel fibers and short tendons, which have been considered particularly well suited for doing positive work on the center of mass, increased their mass-specific energy use during uphill running significantly more than pinnate stance-phase muscles. This finding provides some evidence for a division of labor among muscles used for mechanical work production based on their muscle–tendon architecture. Nevertheless, 33% of the total increase in energy use (40% of the increase in stance-phase energy use) during uphill running was provided by pinnate stance-phase muscles. Swing-phase muscles also increase their energy expenditure during uphill running, although to a lesser extent than that required by running faster on the level. These results suggest that neither muscle–tendon nor musculoskeletal architecture appear to greatly restrict the ability of muscles to do work during locomotor tasks such as uphill running, and that the added energy cost of running uphill is not solely due to lifting the body center of mass.

We examined the energetic cost of loading the trunk or distal portion of the leg in walking and running guinea fowl (Numida meleagris). These different loading regimes were designed to separately influence the energy use by muscles used during the stance and swing phases of the stride. Metabolic rate, estimated from oxygen consumption, was measured while birds locomoted on a motorized treadmill at speeds from 0.5 to 2.0 m s-1, either unloaded, or with a mass equivalent to 23% of their body mass carried on their backs, or with masses equal to approximately 2.5% of their body mass attached to each tarsometatarsal segment. In separate experiments, we also measured the duration of stance and swing in unloaded, trunk-loaded, or limb-loaded birds. In the unloaded and limb-loaded birds, we also calculated the mechanical energy of the tarsometatarsal segment throughout the stride. Trunk and limb loads caused similar increases in metabolic rate. During trunk loading, the net metabolic rate (gross metabolic rate-resting metabolic rate) increased by 17% above the unloaded value across all speeds. This percentage increase is less than has been found in most studies of humans and other mammals. The economical load carriage of guinea fowl is consistent with predictions based on the relative cost of the stance and swing phases of the stride in this species. However, the available comparative data and considerations of the factors that determine the cost of carrying extra mass lead us to the conclusion that the cost of load carrying is unlikely to be a reliable indicator of the distribution of energy use in stance and swing. Both loading regimes caused small changes in the swing and/or stance durations, but these changes were less than 10%. Loading the tarsometatarsal segment increased its segmental energy by 4.1 times and the segmental mechanical power averaged over the stride by 3.8 times. The increases in metabolism associated with limb loading appear to be linked to the increases in mechanical power. The delta efficiency (change in mechanical power divided by the change in metabolic power) of producing this power increased from 11% in walking to approximately 25% in running. Although tarsometatarsal loading was designed to increase the mechanical energy during swing phase, 40% of the increase in segmental energy occurred during late stance. Thus, the increased energy demand of distal limb loading in guinea fowl is predicted to cause increases in energy use by both stance- and swing-phase muscles.