Elastic energy storage and release can enhance performance that would otherwise be limited by the force-velocity constraints of muscle. While 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. At maturity, we compared the jump performance of our treatment group to 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.

ABSTRACT 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; COT JNT ) was maintained at values closer to normal walking than COM mechanical cost of transport (COT COM ; P <0.05, N =15). Furthermore, compared with our measures of COM mechanics (COT COM , COM displacement), joint-level variables (COT JNT , 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.

ABSTRACT 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 ( v walk ) to a variety of running velocities ( v run ; ranging from a sprint to a velocity slower than v walk ). WRTs to fast v run 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 v walk 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 v run , 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 v run 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.

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.

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 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.

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.

SUMMARY The 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 –1 m –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.

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.

SUMMARY Uphill 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.

SUMMARY 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.