Habitually wearing high heels may improve user walking economy in any footwear.

The habitual use of high-heeled footwear may structurally remodel user leg muscle tendons, thereby altering their functional capabilities. High heels set users' ankles in relatively plantarflexed positions, causing calf muscle tendons to operate at relatively short lengths. Habitually operating muscle tendons at relatively short lengths induces structural remodeling that theoretically affects muscle metabolism. Because structural changes occur within the body, the user's locomotor metabolism may change in any footwear condition (e.g., conventional shoes, barefoot). Here, we studied the influence of habitual high-heel use on users' leg muscle-tendon structure and metabolism during walking in flat-soled footwear. We tested eight participants before and after 14 wk of agreeing to wear high heels as their daily shoes. Overall, participants who wore high heels >1,500 steps per day, experienced a 9% decrease in their net metabolic power during walking in flat-soled footwear (d = 1.66, P ≤ 0.049), whereas participants who took <1,000 daily steps in high heels did not (d = 0.44; P = 0.524). Across participants, for every 1,000 daily steps in high heels, net metabolic power during walking in flat-soled footwear decreased 5.3% (r = -0.73; P = 0.040). Metabolic findings were partially explained (r2 = 0.43; P = 0.478) by trending shorter medial gastrocnemius fascicle lengths (d = 0.500, P = 0.327) and increased Achilles tendon stiffness (d = 2.889, P = 0.088). The high-heel intervention did not alter user walking stride kinematics in flat-soled footwear (d ≤ 0.567, P ≥ 0.387). While our limited dataset is unable to establish the mechanisms underlying the high-heel-induced walking economy improvement, it appears that prescribing specific footwear use can be implemented to alter user muscle-tendon properties and augment their function in any shoes.NEW & NOTEWORTHY Habitually wearing high-heeled footwear structurally remodels leg muscle tendons and improves user walking economy, regardless of worn attire.

Correction to: 'Sprinting with prosthetic versus biological legs: insight from experimental data' (2023) by Beck et al.

[This corrects the article DOI: 10.1098/rsos.211799.][This corrects the article DOI: 10.1098/rsos.211799.].

Equivalent running leg lengths require prosthetic legs to be longer than biological legs during standing.

We aimed to determine a method for prescribing a standing prosthetic leg length (ProsL) that results in an equivalent running biological leg length (BioL) for athletes with unilateral (UTTA) and bilateral transtibial amputations (BTTA). We measured standing leg length of ten non-amputee (NA) athletes, ten athletes with UTTA, and five athletes with BTTA. All athletes performed treadmill running trials from 3 m/s to their maximum speed. We calculated standing and running BioL and ProsL lengths and assessed the running-to-standing leg length ratio (Lratio) at three instances during ground contact: touchdown, mid-stance, and take-off. Athletes with UTTA had 2.4 cm longer standing ProsL than BioL length (p = 0.030), but their ProsL length were up to 3.3 cm shorter at touchdown and 4.1 cm shorter at mid-stance than BioL, at speed 3-11.5 m/s. At touchdown, mid-stance, and take-off, athletes with BTTA had 0.01-0.05 lower Lratio at 3 m/s (p < 0.001) and 0.03-0.07 lower Lratio at 10 m/s (p < 0.001) in their ProsL compared to the BioL of NA athletes. During running, ProsL were consistently shorter than BioL. To achieve equivalent running leg lengths at touchdown and take-off, athletes with UTTA should set their running-specific prosthesis height so that their standing ProsL length is 2.8-4.5% longer than their BioL length, and athletes with BTTA should set their running-specific prosthesis height so that their standing ProsL lengths are at least 2.1-3.9% longer than their presumed BioL length. Setting ProsL length to match presumed biological dimensions during standing results in shorter legs during running.

Voluntary muscle coactivation in quiet standing elicits reciprocal rather than coactive agonist-antagonist control of reactive balance.

Muscle coactivation increases in challenging balance conditions as well as with advanced age and mobility impairments. Increased muscle coactivation can occur both in anticipation of (feedforward) and in reaction to (feedback) perturbations, however, the causal relationship between feedforward and feedback muscle coactivation remains elusive. Here, we hypothesized that feedforward muscle coactivation would increase both the body's initial mechanical resistance due to muscle intrinsic properties and the later feedback-mediated muscle coactivation in response to postural perturbations. Young adults voluntarily increased leg muscle coactivation using visual biofeedback before support-surface perturbations. In contrast to our hypothesis, feedforward muscle coactivation did not increase the body's initial intrinsic resistance to perturbations, nor did it increase feedback muscle coactivation. Rather, perturbations with feedforward muscle coactivation elicited a medium- to long-latency increase of feedback-mediated agonist activity but a decrease of feedback-mediated antagonist activity. This reciprocal rather than coactivation effect on ankle agonist and antagonist muscles enabled faster reactive ankle torque generation, reduced ankle dorsiflexion, and reduced center of mass (CoM) motion. We conclude that in young adults, voluntary feedforward muscle coactivation can be independently modulated with respect to feedback-mediated muscle coactivation. Furthermore, our findings suggest feedforward muscle coactivation may be useful for enabling quicker joint torque generation through reciprocal, rather than coactivated, agonist-antagonist feedback muscle activity. As such our results suggest that behavioral context is critical to whether muscle coactivation functions to increase agility versus stability.NEW & NOTEWORTHY Feedforward and feedback muscle coactivation are commonly observed in older and mobility impaired adults and are considered strategies to improve stability by increasing body stiffness prior to and in response to perturbations. In young adults, voluntary feedforward coactivation does not necessarily increase feedback coactivation in response to perturbations. Instead, feedforward coactivation enabled faster ankle torques through reciprocal agonist-antagonist muscle activity. As such, coactivation may promote either agility or stability depending on the behavioral context.

It is time to abandon single-value oxygen uptake energy equivalents.

Physiologists commonly use single-value energy equivalents (e.g., 20.1 kJ/LO2 and 20.9 kJ/LO2) to convert oxygen uptake (V̇o2) to energy, but doing so ignores how the substrate oxidation ratio (carbohydrate:fat) changes across aerobic intensities. Using either 20.1 kJ/LO2 or 20.9 kJ/LO2 can incur systematic errors of up to 7%. In most circumstances, the best approach for estimating energy expenditure is to measure both V̇o2 and V̇co2 and use accurate, species-appropriate stoichiometry. However, there are circumstances when V̇co2 measurements may be unreliable. In those circumstances, we recommend that the research report or compare only V̇o2.NEW & NOTEWORTHY We quantify that the common practice of using single-value oxygen uptake energy equivalents for exercising subjects can incur systematic errors of up to 7%. We argue that such errors can be greatly reduced if researchers measure both V̇o2 and V̇co2 and adopt appropriate stoichiometry equations.

Exoskeletons need to react faster than physiological responses to improve standing balance.

Maintaining balance throughout daily activities is challenging because of the unstable nature of the human body. For instance, a person's delayed reaction times limit their ability to restore balance after disturbances. Wearable exoskeletons have the potential to enhance user balance after a disturbance by reacting faster than physiologically possible. However, "artificially fast" balance-correcting exoskeleton torque may interfere with the user's ensuing physiological responses, consequently hindering the overall reactive balance response. Here, we show that exoskeletons need to react faster than physiological responses to improve standing balance after postural perturbations. Delivering ankle exoskeleton torque before the onset of physiological reactive joint moments improved standing balance by 9%, whereas delaying torque onset to coincide with that of physiological reactive ankle moments did not. In addition, artificially fast exoskeleton torque disrupted the ankle mechanics that generate initial local sensory feedback, but the initial reactive soleus muscle activity was only reduced by 18% versus baseline. More variance of the initial reactive soleus muscle activity was accounted for using delayed and scaled whole-body mechanics [specifically center of mass (CoM) velocity] versus local ankle-or soleus fascicle-mechanics, supporting the notion that reactive muscle activity is commanded to achieve task-level goals, such as maintaining balance. Together, to elicit symbiotic human-exoskeleton balance control, device torque may need to be informed by mechanical estimates of global sensory feedback, such as CoM kinematics, that precede physiological responses.

Evaluating the 'cost of generating force' hypothesis across frequency in human running and hopping.

The volume of active muscle and duration of extensor muscle force well explain the associated metabolic energy expenditure across body mass and velocity during level-ground running and hopping. However, if these parameters fundamentally drive metabolic energy expenditure, then they should pertain to multiple modes of locomotion and provide a simple framework for relating biomechanics to metabolic energy expenditure in bouncing gaits. Therefore, we evaluated the ability of the 'cost of generating force' hypothesis to link biomechanics and metabolic energy expenditure during human running and hopping across step frequencies. We asked participants to run and hop at 85%, 92%, 100%, 108% and 115% of preferred running step frequency. We calculated changes in active muscle volume, duration of force production and metabolic energy expenditure. Overall, as step frequency increased, active muscle volume decreased as a result of postural changes via effective mechanical advantage (EMA) or duty factor. Accounting for changes in EMA and muscle volume better related to metabolic energy expenditure during running and hopping at different step frequencies than assuming a constant EMA and muscle volume. Thus, to ultimately develop muscle mechanics models that can explain metabolic energy expenditure across different modes of locomotion, we suggest more precise measures of muscle force production that include the effects of EMA.

Shorter muscle fascicle operating lengths increase the metabolic cost of cyclic force production.

During locomotion, force-producing limb muscles are predominantly responsible for an animal's whole body metabolic energy expenditure. Animals can change the length of their force-producing muscle fascicles by altering body posture (e.g., joint angles), the structural properties of their biological tissues over time (e.g., tendon stiffness), or the body's kinetics (e.g., body weight). Currently, it is uncertain whether relative muscle fascicle operating lengths have a measurable effect on the metabolic energy expended during cyclic locomotion-like contractions. To address this uncertainty, we quantified the metabolic energy expenditure of human participants, as they cyclically produced two distinct ankle moments at three ankle angles (90°, 105°, and 120°) on a fixed-position dynamometer using their soleus. Overall, increasing participant ankle angle from 90° to 120° (more plantar flexion) reduced minimum soleus fascicle length by 17% (both moment levels, P < 0.001) and increased metabolic energy expenditure by an average of 208% across both moment levels (both P < 0.001). For both moment levels, the increased metabolic energy expenditure was not related to greater fascicle positive mechanical work (higher moment level, P = 0.591), fascicle force rate (both P ≥ 0.235), or model-estimated active muscle volume (both P ≥ 0.122). Alternatively, metabolic energy expenditure correlated with average relative soleus fascicle length (r = -0.72, P = 0.002) and activation (r = 0.51, P < 0.001). Therefore, increasing active muscle fascicle operating lengths may reduce metabolic energy expended during locomotion.NEW & NOTEWORTHY During locomotion, active muscles undergo cyclic length-changing contractions. In this study, we isolated confounding variables and revealed that cyclically producing force at relatively shorter fascicle lengths increases metabolic energy expenditure. Therefore, muscle fascicle operating lengths likely have a measurable effect on the metabolic energy expenditure during locomotion.

Running-specific prosthesis model, stiffness and height affect biomechanics and asymmetry of athletes with unilateral leg amputations across speeds.

Athletes with transtibial amputation (TTA) use running-specific prostheses (RSPs) to run. RSP configuration likely affects the biomechanics of such athletes across speeds. We determined how the use of three RSP models (Catapult, Sprinter and Xtend) with three stiffness categories (recommended, ±1), and three heights (recommended, ±2 cm) affected contact length (Lc ), stance average vertical ground reaction force (F avg), step frequency (f step) and asymmetry between legs for 10 athletes with unilateral TTA at 3-7 m s-1. The use of the Xtend versus Catapult RSP decreased Lc (p = 2.69 × 10-7) and F avg asymmetry (p = 0.032); the effect on Lc asymmetry diminished with faster speeds (p = 0.0020). The use of the Sprinter versus Catapult RSP decreased F avg asymmetry (p = 7.00 × 10-5); this effect was independent of speed (p = 0.90). The use of a stiffer RSP decreased Lc asymmetry (p ≤ 0.00033); this effect was independent of speed (p ≥ 0.071). The use of a shorter RSP decreased Lc (p = 5.86 × 10-6), F avg (p = 8.58 × 10-6) and f step asymmetry (p = 0.0011); each effect was independent of speed (p ≥ 0.15). To minimize asymmetry, athletes with unilateral TTA should use an Xtend or Sprinter RSP with 2 cm shorter than recommended height and stiffness based on intended speed.

Sprinting with prosthetic versus biological legs: insight from experimental data.

Running-prostheses have enabled exceptional athletes with bilateral leg amputations to surpass Olympic 400 m athletics qualifying standards. Due to the world-class performances and relatively fast race finishes of these athletes, many people assume that running-prostheses provide users an unfair advantage over biologically legged competitors during long sprint races. These assumptions have led athletics governing bodies to prohibit the use of running-prostheses in sanctioned non-amputee (NA) competitions, such as at the Olympics. However, here we show that no athlete with bilateral leg amputations using running-prostheses, including the fastest such athlete, exhibits a single 400 m running performance metric that is better than those achieved by NA athletes. Specifically, the best experimentally measured maximum running velocity and sprint endurance profile of athletes with prosthetic legs are similar to, but not better than those of NA athletes. Further, the best experimentally measured initial race acceleration (from 0 to 20 m), maximum velocity around curves, and velocity at aerobic capacity of athletes with prosthetic legs were 40%, 1-3% and 19% slower compared to NA athletes, respectively. Therefore, based on these 400 m performance metrics, use of prosthetic legs during 400 m running races is not unequivocally advantageous compared to the use of biological legs.

Reduced Achilles Tendon Stiffness Disrupts Calf Muscle Neuromechanics in Elderly Gait.

Older adults walk slower and with a higher metabolic energy expenditure than younger adults. In this review, we explore the hypothesis that age-related declines in Achilles tendon stiffness increase the metabolic cost of walking due to less economical calf muscle contractions and increased proximal joint work. This viewpoint may motivate interventions to restore ankle muscle-tendon stiffness, improve walking mechanics, and reduce metabolic cost in older adults.

Adding carbon fiber to shoe soles may not improve running economy: a muscle-level explanation.

In an attempt to improve their distance-running performance, many athletes race with carbon fiber plates embedded in their shoe soles. Accordingly, we sought to establish whether, and if so how, adding carbon fiber plates to shoes soles reduces athlete aerobic energy expenditure during running (improves running economy). We tested 15 athletes as they ran at 3.5 m/s in four footwear conditions that varied in shoe sole bending stiffness, modified by carbon fiber plates. For each condition, we quantified athlete aerobic energy expenditure and performed biomechanical analyses, which included the use of ultrasonography to examine soleus muscle dynamics in vivo. Overall, increased footwear bending stiffness lengthened ground contact time (p = 0.048), but did not affect ankle (p ≥ 0.060), knee (p ≥ 0.128), or hip (p ≥ 0.076) joint angles or moments. Additionally, increased footwear bending stiffness did not affect muscle activity (all seven measured leg muscles (p ≥ 0.146)), soleus active muscle volume (p = 0.538; d = 0.241), or aerobic power (p = 0.458; d = 0.04) during running. Hence, footwear bending stiffness does not appear to alter the volume of aerobic energy consuming muscle in the soleus, or any other leg muscle, during running. Therefore, adding carbon fiber plates to shoe soles slightly alters whole-body and calf muscle biomechanics but may not improve running economy.

Cyclically producing the same average muscle-tendon force with a smaller duty increases metabolic rate.

Ground contact duration and stride frequency each affect muscle metabolism and help scientists link walking and running biomechanics to metabolic energy expenditure. While these parameters are often used independently, the product of ground contact duration and stride frequency (i.e. duty factor) may affect muscle contractile mechanics. Here, we sought to separate the metabolic influence of the duration of active force production, cycle frequency and duty factor. Human participants produced cyclic contractions using their soleus (which has a relatively homogeneous fibre type composition) at prescribed cycle-average ankle moments on a fixed dynamometer. Participants produced these ankle moments over short, medium and long durations while maintaining a constant cycle frequency. Overall, decreased duty factor did not affect cycle-average fascicle force (p ≥ 0.252) but did increase net metabolic power (p ≤ 0.022). Mechanistically, smaller duty factors increased maximum muscle-tendon force (p < 0.001), further stretching in-series tendons and shifting soleus fascicles to shorter lengths and faster velocities, thereby increasing soleus total active muscle volume (p < 0.001). Participant soleus total active muscle volume well-explained net metabolic power (r = 0.845; p < 0.001). Therefore, cyclically producing the same cycle-average muscle-tendon force using a decreased duty factor increases metabolic energy expenditure by eliciting less economical muscle contractile mechanics.

Added lower limb mass does not affect biomechanical asymmetry but increases metabolic power in runners with a unilateral transtibial amputation.

PURPOSE: we determined the metabolic and biomechanical effects of adding mass to the running-specific prosthesis (RSP) and biological foot of individuals with a unilateral transtibial amputation (TTA) during running. METHODS: 10 individuals (8 males, 2 females) with a TTA ran on a force-measuring treadmill at 2.5 m/s with 100 g and 300 g added to their RSP alone or to their RSP and biological foot while we measured their metabolic rates and calculated peak vertical ground reaction force (vGRF), stance-average vGRF, and step time symmetry indices. RESULTS: for every 100 g added to the RSP alone, metabolic power increased by 0.86% (p = 0.007) and for every 100 g added to the RSP and biological foot, metabolic power increased by 1.74% ([Formula: see text] 0.001) during running. Adding mass had no effect on peak vGRF (p = 0.102), stance-average vGRF (p = 0.675), or step time (p = 0.413) symmetry indices. We also found that the swing time of the affected leg was shorter than the unaffected leg across conditions ([Formula: see text] 0.007). CONCLUSIONS: adding mass to the lower limbs of runners with a TTA increased metabolic power by more than what has been reported for those without an amputation. We found no effect of added mass on biomechanical asymmetry, but the affected leg had consistently shorter swing times than the unaffected leg. This suggests that individuals with a TTA maintain asymmetries despite changes in RSP mass and that lightweight prostheses could improve performance by minimizing metabolic power without affecting asymmetry.

The exoskeleton expansion: improving walking and running economy.

Since the early 2000s, researchers have been trying to develop lower-limb exoskeletons that augment human mobility by reducing the metabolic cost of walking and running versus without a device. In 2013, researchers finally broke this 'metabolic cost barrier'. We analyzed the literature through December 2019, and identified 23 studies that demonstrate exoskeleton designs that improved human walking and running economy beyond capable without a device. Here, we reviewed these studies and highlighted key innovations and techniques that enabled these devices to surpass the metabolic cost barrier and steadily improve user walking and running economy from 2013 to nearly 2020. These studies include, physiologically-informed targeting of lower-limb joints; use of off-board actuators to rapidly prototype exoskeleton controllers; mechatronic designs of both active and passive systems; and a renewed focus on human-exoskeleton interface design. Lastly, we highlight emerging trends that we anticipate will further augment wearable-device performance and pose the next grand challenges facing exoskeleton technology for augmenting human mobility.

Prosthetic shape, but not stiffness or height, affects the maximum speed of sprinters with bilateral transtibial amputations.

Running-specific prostheses (RSPs) have facilitated an athlete with bilateral transtibial amputations to compete in the Olympic Games. However, the performance effects of using RSPs compared to biological legs remains controversial. Further, the use of different prosthetic configurations such as shape, stiffness, and height likely influence performance. We determined the effects of using 15 different RSP configurations on the maximum speed of five male athletes with bilateral transtibial amputations. These athletes performed sets of running trials up to maximum speed using three different RSP models (Freedom Innovations Catapult FX6, Össur Flex-Foot Cheetah Xtend and Ottobock 1E90 Sprinter) each with five combinations of stiffness category and height. We measured ground reaction forces during each maximum speed trial to determine the biomechanical parameters associated with different RSP configurations and maximum sprinting speeds. Use of the J-shaped Cheetah Xtend and 1E90 Sprinter RSPs resulted in 8.3% and 8.0% (p<0.001) faster maximum speeds compared to the use of the C-shaped Catapult FX6 RSPs, respectively. Neither RSP stiffness expressed as a category (p = 0.836) nor as kN·m-1 (p = 0.916) affected maximum speed. Further, prosthetic height had no effect on maximum speed (p = 0.762). Faster maximum speeds were associated with reduced ground contact time, aerial time, and overall leg stiffness, as well as with greater stance-average vertical ground reaction force, contact length, and vertical stiffness (p = 0.015 for aerial time, p<0.001 for all other variables). RSP shape, but not stiffness or height, influences the maximum speed of athletes with bilateral transtibial amputations.

Prosthetic model, but not stiffness or height, affects maximum running velocity in athletes with unilateral transtibial amputations.

The running-specific prosthetic (RSP) configuration used by athletes with transtibial amputations (TTAs) likely affects performance. Athletes with unilateral TTAs are prescribed C- or J-shaped RSPs with a manufacturer-recommended stiffness category based on body mass and activity level, and height based on unaffected leg and residual limb length. We determined how 15 different RSP model, stiffness, and height configurations affect maximum running velocity (vmax) and the underlying biomechanics. Ten athletes with unilateral TTAs ran at 3 m/s to vmax on a force-measuring treadmill. vmax was 3.8-10.7% faster when athletes used J-shaped versus C-shaped RSP models (p < 0.05), but was not affected by stiffness category, actual stiffness (kN/m), or height (p = 0.72, p = 0.37, and p = 0.11, respectively). vmax differences were explained by vertical ground reaction forces (vGRFs), stride kinematics, leg stiffness, and symmetry. While controlling for velocity, use of J-shaped versus C-shaped RSPs resulted in greater stance average vGRFs, slower step frequencies, and longer step lengths (p < 0.05). Stance average vGRFs were less asymmetric using J-shaped versus C-shaped RSPs (p < 0.05). Contact time and leg stiffness were more asymmetric using the RSP model that elicited the fastest vmax (p < 0.05). Thus, RSP geometry (J-shape versus C-shape), but not stiffness or height, affects vmax in athletes with unilateral TTAs.

Exoskeletons Improve Locomotion Economy by Reducing Active Muscle Volume.

Exoskeletons that improve locomotion economy typically are engineered to reduce users' limb joint mechanical work or moments. Yet, limb joint dynamics do not necessarily reflect muscle dynamics, which dictate whole-body metabolic energy expenditure. Here, we hypothesize that exoskeletons primarily reduce user metabolic energy expenditure across locomotion conditions by reducing active muscle volume.