Metabolic cost of generating horizontal forces during human running.

Previous studies have suggested that generating vertical force on the ground to support body weight (BWt) is the major determinant of the metabolic cost of running. Because horizontal forces exerted on the ground are often an order of magnitude smaller than vertical forces, some have reasoned that they have negligible cost. Using applied horizontal forces (AHF; negative is impeding, positive is aiding) equal to -6, -3, 0, +3, +6, +9, +12, and +15% of BWt, we estimated the cost of generating horizontal forces while subjects were running at 3.3 m/s. We measured rates of oxygen consumption (VO2) for eight subjects. We then used a force-measuring treadmill to measure ground reaction forces from another eight subjects. With an AHF of -6% BWt, VO2 increased 30% compared with normal running, presumably because of the extra work involved. With an AHF of +15% BWt, the subjects exerted approximately 70% less propulsive impulse and exhibited a 33% reduction in VO2. Our data suggest that generating horizontal propulsive forces constitutes more than one-third of the total metabolic cost of normal running.     

Energetics of walking and running: insights from simulated reduced-gravity experiments.

On Earth, a person uses about one-half as much energy to walk a mile as to run a mile. On another planet with lower gravity, would walking still be more economical than running? When people carry weights while they walk or run, energetic cost increases in proportion to the added load. It would seem to follow that if gravity were reduced, energetic cost would decrease in proportion to body weight in both gaits. However, we find that under simulated reduced gravity, the rate of energy consumption decreases in proportion to body weight during running but not during walking. When gravity is reduced by 75%, the rate of energy consumption is reduced by 72% during running but only by 33% during walking. Because reducing gravity decreases the energetic cost much more for running than for walking, walking is not the cheapest way to travel a mile at low levels of gravity. These results suggest that the link between the mechanics of locomotion and energetic cost is fundamentally different for walking and for running.     

Metabolic cost of walking: equation and model

Twenty years of published experience with the Workman-Armstrong equation for predicting walking VO2 is reviewed. The equation is reexpressed in currently accepted terminology, and it is shown that the equation serves well as a basic model of normal walking. Employing this model to analyze VO2/step leads to the elaboration of a three-compartment model of the metabolic cost of walking. This three-compartment model provides a rational estimate of the fraction of walking's metabolic cost that powers the actual walking movement. Doubt is expressed that "comfortable speed of walking" is definable in energy terms. It is suggested that the requirements of maintaining balance while walking may determine both the comfortable speed of walking and the curvilinearity of the relationship between ground-speed and freely chosen step frequency of walking.     

Energy cost and muscular activity required for propulsion during walking.

We reasoned that with an optimal aiding horizontal force, the reduction in metabolic rate would reflect the cost of generating propulsive forces during normal walking. Furthermore, the reductions in ankle extensor electromyographic (EMG) activity would indicate the propulsive muscle actions. We applied horizontal forces at the waist, ranging from 15% body weight aiding to 15% body weight impeding, while subjects walked at 1.25 m/s. With an aiding horizontal force of 10% body weight, 1) the net metabolic cost of walking decreased to a minimum of 53% of normal walking, 2) the mean EMG of the medial gastrocnemius (MG) during the propulsive phase decreased to 59% of the normal walking magnitude, and yet 3) the mean EMG of the soleus (Sol) did not decrease significantly. Our data indicate that generating horizontal propulsive forces constitutes nearly half of the metabolic cost of normal walking. Additionally, it appears that the MG plays an important role in forward propulsion, whereas the Sol does not.     

Energy cost and muscular activity required for leg swing during walking.

To investigate the metabolic cost and muscular actions required for the initiation and propagation of leg swing, we applied a novel combination of external forces to subjects walking on a treadmill. We applied a forward pulling force at each foot to assist leg swing, a constant forward pulling force at the waist to provide center of mass propulsion, and a combination of these foot and waist forces to evaluate leg swing. When the metabolic cost and muscle actions were at a minimum, the condition was considered optimal. We reasoned that the difference in energy consumption between the optimal combined waist and foot force trial and the optimal waist force-only trial would reflect the metabolic cost of initiating and propagating leg swing during normal walking. We also reasoned that a lower muscle activity with these assisting forces would indicate which muscles are normally responsible for initiating and propagating leg swing. With a propulsive force at the waist of 10% body weight (BW), the net metabolic cost of walking decreased to 58% of normal walking. With the optimal combination, a propulsive force at the waist of 10% BW plus a pulling force at the feet of 3% BW the net metabolic cost of walking further decreased to 48% of normal walking. With the same combination, the muscle activity of the iliopsoas and rectus femoris muscles during the swing phase was 27 and 60% lower, respectively, but the activity of the medial gastrocnemius and soleus before swing did not change. Thus our data indicate that approximately 10% of the net metabolic cost of walking is required to initiate and propagate leg swing. Additionally, the hip flexor muscles contribute to the initiation and propagation leg swing.     

Effects of obesity and sex on the energetic cost and preferred speed of walking.

The metabolic energy cost of walking is determined, to a large degree, by body mass, but it is not clear how body composition and mass distribution influence this cost. We tested the hypothesis that walking would be most expensive for obese women compared with obese men and normal-weight women and men. Furthermore, we hypothesized that for all groups, preferred walking speed would correspond to the speed that minimized the gross energy cost per distance. We measured body composition, maximal oxygen consumption, and preferred walking speed of 39 (19 class II obese, 20 normal weight) women and men. We also measured oxygen consumption and carbon dioxide production while the subjects walked on a level treadmill at six speeds (0.50-1.75 m/s). Both obesity and sex affected the net metabolic rate (W/kg) of walking. Net metabolic rates of obese subjects were only approximately 10% greater (per kg) than for normal-weight subjects, and net metabolic rates for women were approximately 10% greater than for men. The increase in net metabolic rate at faster walking speeds was greatest in obese women compared with the other groups. Preferred walking speed was not different across groups (1.42 m/s) and was near the speed that minimized gross energy cost per distance. Surprisingly, mass distribution (thigh mass/body mass) was not related to net metabolic rate, but body composition (% fat) was (r2= 0.43). Detailed biomechanical studies of walking are needed to investigate whether obese individuals adopt novel energy saving mechanisms during walking.     

Muscle mechanical advantage of human walking and running: implications for energy cost.

Muscular forces generated during locomotion depend on an animal's speed, gait, and size and underlie the energy demand to power locomotion. Changes in limb posture affect muscle forces by altering the mechanical advantage of the ground reaction force (R) and therefore the effective mechanical advantage (EMA = r/R, where r is the muscle mechanical advantage) for muscle force production. We used inverse dynamics based on force plate and kinematic recordings of humans as they walked and ran at steady speeds to examine how changes in muscle EMA affect muscle force-generating requirements at these gaits. We found a 68% decrease in knee extensor EMA when humans changed gait from a walk to a run compared with an 18% increase in hip extensor EMA and a 23% increase in ankle extensor EMA. Whereas the knee joint was extended (154-176 degrees) during much of the support phase of walking, its flexed position (134-164 degrees) during running resulted in a 5.2-fold increase in quadriceps impulse (time-integrated force during stance) needed to support body weight on the ground. This increase was associated with a 4.9-fold increase in the ground reaction force moment about the knee. In contrast, extensor impulse decreased 37% (P < 0.05) at the hip and did not change at the ankle when subjects switched from a walk to a run. We conclude that the decrease in limb mechanical advantage (mean limb extensor EMA) and increase in knee extensor impulse during running likely contribute to the higher metabolic cost of transport in running than in walking. The low mechanical advantage in running humans may also explain previous observations of a greater metabolic cost of transport for running humans compared with trotting and galloping quadrupeds of similar size.     

The independent effects of speed and propulsive force on joint power generation in walking

Walking speed is modulated using propulsive forces (F_P) during push-off and both preferred speed and F_P decrease with aging. However, even prior to walking slower, reduced F_P may be accompanied by potentially unfavorable changes in joint power generation. For example, compared to young adults, older adults exhibit a redistribution of mechanical power generation from the propulsive plantarflexor muscles to more proximal muscles acting across the knee and hip. Here, we used visual biofeedback based on real-time F_P measurements to decouple and investigate the interaction between joint-level coordination, whole-body F_P, and walking speed. 12 healthy young subjects walked on a dual-belt instrumented treadmill at a range of speeds (0.9–1.3 m/s). We immediately calculated the average F_P from each speed. Subjects then walked at 1.3 m/s while completing a series of biofeedback trials with instructions to match their instantaneous F_P to their averaged F_P from slower speeds. Walking slower decreased F_P and total positive joint work with little effect on relative joint-level contributions. Conversely, subjects walked at a constant speed with reduced F_P, not by reducing total positive joint work, but by redistributing the mechanical demands of each step from the plantarflexor muscles during push-off to more proximal leg muscles during single support. Interestingly, these naturally emergent joint- and limb-level biomechanical changes, in the absence of neuromuscular constraints, resemble those due to aging. Our findings provide important reference data to understand the presumably complex interactions between joint power generation, whole-body F_P, and walking speed in our aging population.     

Metabolic cost of motility in planktonic protists: Theoretical considerations on size scaling and swimming speed

The metabolic cost of swimming for planktonic protists is calculated, on theoretical grounds, from a simple model based upon Stokes' law. Energetic expenditure is scaled over both typically encountered size ranges (1–100 µm) and swimming speeds (100–5,000 µm/sec). In agreement with previous estimates for typical flagellates, these estimates generally suggest a low (<1%) cost for motility, related to total metabolic rate of growing cells. However, the cost of motility in small, fast-moving forms, such as some ciliates and flagellates, may be significant (1–10%) and even substantial (10–100%+) for certain species. In accordance with these predictions, many fast-moving ciliates restrict motility to bursts of activity or jumps. In the absence of a reduction in swimming speed or in the frequency of jumps, it is predicted that this relative cost of motility will be significantly increased in starving heterotrophs or light-limited autotrophs, if such cells reduce cell volumes and specific rates of respiration.     

Predicting the metabolic cost of incline walking from muscle activity and walking mechanics

The goal of this study was to identify which muscle activation patterns and gait features best predict the metabolic cost of inclined walking. We measured muscle activation patterns, joint kinematics and kinetics, and metabolic cost in sixteen subjects during treadmill walking at inclines of 0%, 5%, and 10%. Multivariate regression models were developed to predict the net metabolic cost from selected groups of the measured variables. A linear regression model including incline and the squared integrated electromyographic signals of the soleus and vastus lateralis explained 96% of the variance in metabolic cost, suggesting that the activation patterns of these large muscles have a high predictive value for metabolic cost. A regression model including only the peak knee flexion angle during stance phase, peak knee extension moment, peak ankle plantarflexion moment, and peak hip flexion moment explained 89% of the variance in metabolic cost; this finding indicates that kinematics and kinetics alone can predict metabolic cost during incline walking. The ability of these models to predict metabolic cost from muscle activation patterns and gait features points the way toward future work aimed at predicting metabolic cost when gait is altered by changes in neuromuscular control or the use of an assistive technology.     

The effects of sensory loss and walking speed on the orbital dynamic stability of human walking

Peripheral sensory feedback is believed to contribute significantly to maintaining walking stability. Patients with diabetic peripheral neuropathy have a greatly increased risk of falling. Previously, we demonstrated that slower walking speeds in neuropathic patients lead to improved local dynamic stability. However, all subjects exhibited significant local instability during walking, even though no subject fell or stumbled during testing. The present study was conducted to determine if and how significant changes in peripheral sensation and walking speed affect orbital stability during walking. Trunk and lower extremity kinematics were examined from two prior experiments that compared patients with significant neuropathy to healthy controls and walking at multiple different speeds in young healthy subjects. Maximum Floquet multipliers were computed for each time series to quantify the orbital stability of these movements. All subjects exhibited orbitally stable walking kinematics, even though these same kinematics were previously shown to be locally unstable. Differences in orbital stability between neuropathic and control subjects were small and, with the exception of knee joint movements (p=0.001), not statistically significant (0.380p0.946). Differences in knee orbital stability were not mediated by differences in walking speed. This was supported by our finding that although orbital stability improved slightly with slower walking speeds, the correlations between walking speed and orbital stability were generally weak (r(2)16.7%). Thus, neuropathic patients do not gain improved orbital stability as a result of slowing down and do not experience any loss of orbital stability because of their sensory deficits.     

More than energy cost: multiple benefits of the long Achilles tendon in human walking and running

Elastic strain energy that is stored and released from long, distal tendons such as the Achilles during locomotion allows for muscle power amplification as well as for reduction of the locomotor energy cost: as distal tendons perform mechanical work during recoil, plantar flexor muscle fibres can work over smaller length ranges, at slower shortening speeds, and at lower activation levels. Scant evidence exists that long distal tendons evolved in humans (or were retained from our more distant Hominoidea ancestors) primarily to allow high muscle–tendon power outputs, and indeed we remain relatively powerless compared to many other species. Instead, the majority of evidence suggests that such tendons evolved to reduce total locomotor energy cost. However, numerous additional, often unrecognised, advantages of long tendons may speculatively be of greater evolutionary advantage, including the reduced limb inertia afforded by shorter and lighter muscles (reducing proximal muscle force requirement), reduced energy dissipation during the foot–ground collisions, capacity to store and reuse the muscle work done to dampen the vibrations triggered by foot–ground collisions, reduced muscle heat production (and thus core temperature), and attenuation of work-induced muscle damage. Cumulatively, these effects should reduce both neuromotor fatigue and sense of locomotor effort, allowing humans to choose to move at faster speeds for longer. As these benefits are greater at faster locomotor speeds, they are consistent with the hypothesis that running gaits used by our ancestors may have exerted substantial evolutionary pressure on Achilles tendon length. The long Achilles tendon may therefore be a singular adaptation that provided numerous physiological, biomechanical, and psychological benefits and thus influenced behaviour across multiple tasks, both including and additional to locomotion. While energy cost may be a variable of interest in locomotor studies, future research should consider the broader range of factors influencing our movement capacity, including our decision to move over given distances at specific speeds, in order to understand more fully the effects of Achilles tendon function as well as changes in this function in response to physical activity, inactivity, disuse and disease, on movement performance.     

Accidental experiments: ecological and evolutionary insights and opportunities derived from global change

Humans are the dominant ecological and evolutionary force on the planet today, transforming habitats, polluting environments, changing climates, introducing new species, and causing other species to decline in number or go extinct. These worrying anthropogenic impacts, collectively termed global change, are often viewed as a confounding factor to minimize in basic studies and a problem to resolve or quantify in applied studies. However, these ‘accidental experiments’ also represent opportunities to gain fundamental insight into ecological and evolutionary processes, especially when they result in perturbations that are large or long in duration and difficult or unethical to impose experimentally. We demonstrate this by describing important fundamental insights already gained from studies which utilize global change factors as accidental experiments. In doing so, we highlight why accidental experiments are sometimes more likely to yield insights than traditional approaches. Next, we argue that emerging environmental problems can provide even more opportunities for scientific discovery in the future, and provide both examples and guidelines for moving forward. We recommend 1) a greater flow of information between basic and applied subfields of ecology and evolution to identify emerging opportunities; 2) considering the advantages of the ‘accidental experiment’ approach relative to more traditional approaches; and 3) planning for the challenges inherent to uncontrolled accidental experiments. We emphasize that we do not view the accidental experiments provided by global change as replacements for scientific studies quantifying the magnitude of anthropogenic impacts or outlining strategies for mitigating impacts. Instead, we believe that accidental experiments are uniquely situated to provide insights into evolutionary and ecological processes that ultimately allow us to better predict and manage change on our human‐dominated planet. Synthesis Humans have an increasingly large impact on the planet. In response, ecologists and evolutionary biologists are dedicating increasing scientific attention to global change, largely with studies documenting biological effects and testing strategies to avoid or reverse negative impacts. In this article, we analyze global change from a different perspective, and suggest that human impacts on the environment also serve as valuable ‘accidental experiments’ that can provide fundamental scientific insight. We highlight and synthesize examples of studies taking this approach, and give guidance for gaining future insights from these unfortunate ‘accidental experiments’.     

Speed training with body weight unloading improves walking energy cost and maximal speed in 75- to 85-year-old healthy women.

This randomized controlled study was designed to prove the hypothesis that a novel approach to high-speed interval training, based on walking on a treadmill with the use of body weight unloading (BWU), would have improved energy cost and speed of overground walking in healthy older women. Participants were randomly assigned to either the exercise group (n = 11, 79.6 +/- 3.7 yr, mean +/- SD) or the nonintervention control group (n = 11, 77.6 +/- 2.3 yr). During the first 6 wk, the exercise group performed walking interval training on the treadmill with 40% BWU at the maximal walking speed corresponding to an intensity close to heart rate at ventilatory threshold (T(vent) walking speed). Each session consisted of four sets of 5 min of walking (three 1-min periods at T(vent) walking speed, with two 1-min intervals at comfortable walking speed in between each period at T(vent) walking speed) with 1-min interval between each set. Speed was increased session by session until the end of week 6. BWU was then progressively reduced to 10% during the last 6 wk of intervention. After 12 wk, the walking energy cost per unit of distance at all self-selected overground walking speeds (slow, comfortable, and fast) was significantly reduced in the range from 18 to 21%. The exercise group showed a 13% increase in maximal walking speed and a 67% increase in mechanical power output at T(vent) after the training program. The novel "overspeed" training approach has been demonstrated to be effective in improving energy cost and speed of overground walking in healthy older women.     

The cost of defense in social insects: insights from the honey bee

Defense is one of the most important factors affecting life history. The relationship of defense to life history traits as well as its possible costs has been reviewed extensively for many groups, including plants. However, defense in social insects, such as honey bees, has never been examined from a trade‐off perspective, although defense in honey bees, Apis mellifera L. (Hymenoptera: Apidae), has been widely studied. In this review, we discuss the life history traits of honey bees, particularly traits related to defense. We then examine trade‐offs in the context of resource availability. Lastly, we offer suggestions for future research on trade‐offs in honey bees and other social insects.     

Effects of aging and arm swing on the metabolic cost of stability in human walking

To gain insight into the mechanical determinants of walking energetics, we investigated the effects of aging and arm swing on the metabolic cost of stabilization. We tested two hypotheses: (1) elderly adults consume more metabolic energy during walking than young adults because they consume more metabolic energy for lateral stabilization, and (2) arm swing reduces the metabolic cost of stabilization during walking in young and elderly adults. To test these hypotheses, we provided external lateral stabilization by applying bilateral forces (10% body weight) to a waist belt via elastic cords while young and elderly subjects walked at 1.3m/s on a motorized treadmill with arm swing and with no arm swing. We found that the external stabilizer reduced the net rate of metabolic energy consumption to a similar extent in elderly and young subjects. This reduction was greater (6-7%) when subjects walked with no arm swing than when they walked normally (3-4%). When young or elderly subjects eliminated arm swing while walking with no external stabilization, net metabolic power increased by 5-6%. We conclude that the greater metabolic cost of walking in elderly adults is not caused by a greater cost of lateral stabilization. Moreover, arm swing reduces the metabolic cost of walking in both young and elderly adults likely by contributing to stability.     

The cost of myrmecophytism: insights from allometry of stem secondary growth

Background and Aims

Plant defence traits against herbivores incur production costs that are usually difficult to measure. However, estimating these costs is a prerequisite for characterizing the plant defence strategy as a whole. Myrmecophytes are plants that provide symbiotic ants with specialized nesting cavities, called domatia, in exchange for protection against herbivores. In the particular case of stem domatia, production of extra wood seems to be the only associated cost, making this indirect defence trait a particularly suitable model for estimating the cost of defence.

Methods

Measurements were made of growth pattern and cumulative production cost of domatia over secondary growth in the myrmecophyte Leonardoxa africana subsp. africana, whose internodes display both a solid basal segment and a hollow distal part (the domatium), thus allowing paired comparison of investment in wood.

Key Results

Previous studies showed that ‘overconstruction’ of the hollow part of internodes during primary growth is needed for mechanical support. In this study, it is shown that the relationship between the woody cross-sectional area of the solid and hollow parts of internodes is negatively allometric at the beginning of secondary growth and nearly isometric later on. Thus, in hollow stems, the first phase of slow secondary growth compensates for the ‘overconstruction’ of the ring of wood during primary growth. Moreover, the cumulative production cost of a domatium (estimated as the additional volume of wood required for a hollow stem compared with a solid one) is very high at the beginning of secondary growth and then quickly tends to zero.

Conclusions

Making domatia incurs high costs early in ontogeny, costs that are then amortized later in development of stems and of individual plants. Characterizing ontogenetic variation of the net cost of this peculiar defence mechanism will help us build more accurate theoretical models of resource allocation in myrmecophytes.