Better economy in field running than on the treadmill: evidence from high-level distance runners

Given the ongoing interest in ways to improve the specificity of testing elite athletes in their natural environment, portable metabolic systems provide an opportunity to assess metabolic demand of exercise in sport-specific settings. Running economy (RE) and maximal oxygen uptake (V.O₂max) were compared between track and treadmill (1% inclination) conditions in competitive level European distance runners who were fully habituated to treadmill running (n = 13). All runners performed an exercise test on running track and on treadmill. While V.O₂max was similar on the track and on the treadmill (68.5 ± 5.3 vs. 71.4 ± 6.4 ml·kg⁻¹·min⁻¹, p = 0.105, respectively), superior RE was found on the track compared to the treadmill (215.4 ± 12.4 vs. 236.8 ± 18.0 O₂ ml·kg⁻¹·km⁻¹, p < 0.001). RE on the track was strongly correlated with RE on the treadmill (r = 0.719, p = 0.006). The present findings indicate that high-level distance runners have significantly better RE but not V.O₂max on the track compared to treadmill. This difference may be due to biomechanical adjustments. As RE is strongly correlated between the two conditions, it would be reasonable to assume that interventions affecting RE on the treadmill will also affect RE on the track.     

Old men running: mechanical work and elastic bounce

It is known that muscular force is reduced in old age. We investigate what are the effects of this phenomenon on the mechanics of running. We hypothesized that the deficit in force would result in a lower push, causing reduced amplitude of the vertical oscillation, with smaller elastic energy storage and increased step frequency. To test this hypothesis, we measured the mechanical energy of the centre of mass of the body during running in old and young subjects. The amplitude of the oscillation is indeed reduced in the old subjects, resulting in an approximately 20% smaller elastic recovery and a greater step frequency (3.7 versus 2.8 Hz, p=1.9x10(-5), at 15-17 km h(-1)). Interestingly, the greater step frequency is due to a lower aerial time, and not to a greater natural frequency of the system, which is similar in old and young subjects (3.6 versus 3.4 Hz, p=0.2). Moreover, we find that in the old subjects, the step frequency is always similar to the natural frequency, even at the highest speeds. This is at variance with young subjects who adopt a step frequency lower than the natural frequency at high speeds, to contain the aerobic energy expenditure. Finally, the external work to maintain the motion of the centre of mass is reduced in the old subjects (0.9 versus 1.2 J kg(-1) m(-1), p=5.1x10(-6)) due to the lower work done against gravity, but the higher step frequency involves a greater internal work to reset the limbs at each step. The net result is that the total work increases with speed more steeply in the old subjects than in young subjects.     

Shaking a leg and hot to trot: the effects of body size and temperature on running speed in ants

Abstract.  1. Data were compiled from the literature and our own studies on 24 ant species to characterise the effects of body size and temperature on forager running speed.
2. Running speed increases with temperature in a manner consistent with the effects of temperature on metabolic rate and the kinetic properties of muscles.
3. The exponent of the body mass-running speed allometry ranged from 0.14 to 0.34 with a central tendency of approximately 0.25. This body mass scaling is consistent with both the model of elastic similarity, and a model combining dynamic similarity with available metabolic power.
4. Even after controlling for body size or temperature, a substantial amount of inter-specific variation in running speed remains. Species with certain lifestyles [e.g. nomadic group predators, species which forage at extreme (>60 °C) temperatures] may have been selected for faster running speeds.
5. Although ants have a similar scaling exponent to mammals for the running speed allometry, they run slower than predicted compared with a hypothetical mammal of similar size. This may in part reflect physiological differences between invertebrates and vertebrates.     

Influence of the oxygen uptake slow component on the aerobic energy cost of high-intensity submaximal treadmill running in humans

During high-intensity running, the oxygen uptake (V˙O₂) kinetics is characterised by a slow component which delays the attainment of the steady-state beyond the 3rd min of exercise. To assess if the aerobic energy cost of running measured at the 3rd min (C ₃) adequately reflects the variability of the true aerobic energy cost measured during the steady-state (C _ss), 13 highly-trained runners completed sessions of square-wave running at intensities above 80% maximal oxygen uptake (V˙O_2max) on a level treadmill. To evaluate the time at which the steady-state V˙O₂ was attained (t _ss), the V˙O₂ responses were described using a general double-exponential equation and t _ss was defined as the time at which V˙O₂ was less than 1% below the asymptotic value given by the model. All the subjects achieved a steady state for intensities equal to or greater than 92% V˙O_2max, and 8 out of 13 achieved it at 99% V˙O_2max. In all cases, t _ss was less than 13 min. For intensities greater than 85% V˙O_2max, C _ss was significantly higher than C ₃ and was positively related to %V˙O_2max (r= 0.44; P < 0.001) while C ₃ remained constant. The C ₃ only explained moderately the variability of C _ss (0.39 < r ² < 0.72, depending on the velocity or the (relative intensity at which the relationship was calculated). Moreover, the excess aerobic energy cost of running the (difference between C _ss and C ₃) was well predicted by age (0.90 < r ² < 0.93). Therefore, when the aerobic profile of runners is evaluated, it is recommended that their running efficiencies at velocities which reflect their race intensities should be determined, with V˙O₂ data being measured at the true steady-state. Accepted: 1 June 1998     

The relationship between limb morphology, kinematics, and force during running: the evolution of locomotor dynamics in lizards

Terrestrial locomotion occurs via the hierarchical links between morphology, kinematics, force, and center-of-mass mechanics. In a phylogenetically broad sample of seven lizard species, we show that morphological variation drives kinematic variation, which, in turn, drives force variation. Species with short limbs use a short stride–high frequency strategy when running at steady-speed and to change speeds. This link between morphology and kinematics results in relatively small vertical forces during the support phase of the stride cycle. Conversely, species with long limbs use a long stride–low frequency strategy, resulting in large vertical forces during the support phase. In view of these findings, we suggest that limb length may predict locomotor energetics in lizards because energetics are largely determined by vertical forces and stride frequency. Additionally, we propose an energetic trade-off with both long- and short-limbed species paying the most energy to move, whereas intermediate-limbed species move using less energy. Finally, when these traits are mapped onto a lizard phylogeny, we show that locomotor functional morphology exhibits both deep phylogenetic effects and contemporary patterns of evolutionary convergence. Overall, the present study provides a foundation for testing hypotheses regarding the integration and evolution of functional traits in lizards and animals in general.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 634–651.     

Electromyographic correlates of the transition from aerobic to anaerobic metabolism in treadmill running

This study analysed the changes in electromyographic (EMG) activity of the vastus lateralis, biceps femoris and gastrocnemius muscles during incremental treadmill running. The changes in EMG were related to the lactate and ventilatory thresholds. Ten trained subjects participated in the study. Minute ventilation, oxygen consumption, carbon dioxide expired and the fraction of oxygen in the expired gas were recorded continuously. Venous blood samples were collected at each exercise intensity and analysed for lactate concentration. The EMG were recorded at the end of each exercise intensity using surface electrodes. The EMG were quantified through integration (iEMG) and by calculating the mean power frequency (MPF). The iEMG measurements were characterized by a breakpoint in the vastus lateralis and/or gastrocnemius muscles in eight of the subjects tested. However, the results indicated that blood lactate concentrations had already begun to increase in a nonlinear fashion before the iEMG breakpoint had been surpassed. Consequently, the occurence of the lactate threshold cannot be attributed solely to the change in motor unit recruitment or rate coding patterns demonstrated by the iEMG breakpoint. The ventilatory threshold was shown to be a far more reliable and convenient noninvasive predictor of the lactate threshold in comparison with EMG techniques. In conclusion, the EMG measurements used in this study (i.e. iEMG and MPF) were not considered to be viable noninvasive determinants of the aerobic-anaerobic transition phase in treadmill running.     

Determinants of gait stability while walking on a treadmill: A machine learning approach

Dynamic balance in human locomotion can be assessed through the local dynamic stability (LDS) method. Whereas gait LDS has been used successfully in many settings and applications, little is known about its sensitivity to individual characteristics of healthy adults. Therefore, we reanalyzed a large dataset of accelerometric data measured for 100 healthy adults from 20 to 70 years of age performing 10 min treadmill walking. We sought to assess the extent to which the variations of age, body mass and height, sex, and preferred walking speed (PWS) could influence gait LDS. The random forest (RF) and multiple adaptive regression splines (MARS) algorithms were selected for their good bias-variance tradeoff and their capabilities to handle nonlinear associations. First, through variable importance measure (VIM), we used RF to evaluate which individual characteristics had the highest influence on gait LDS. Second, we used MARS to detect potential interactions among individual characteristics that may influence LDS. The VIM and MARS results indicated that PWS and age correlated with LDS, whereas no associations were found for sex, body height, and body mass. Further, the MARS model detected an age by PWS interaction: on one hand, at high PWS, gait stability is constant across age while, on the other hand, at low PWS, gait instability increases substantially with age. We conclude that it is advisable to consider the participants’ age as well as their PWS to avoid potential biases in evaluating dynamic balance through LDS.     

Squirrel monkey locomotion on an inclined treadmill: Implications for the evolution of gaits

Three adult squirrel monkeys were trained to run on a motor-driven treadmill that was inclined downwardly and upwardly at 8°, 16° and 28°, and horizontally (0°). Films were used to compare the gait and kinematics of the animals across the inclines. All three animals used both lateral and diagonal sequence gaits, although the former was preferred at all but the upward 16° and 28° inclines. Cycle duration and hind limb stance and swing durations tended to increase as downward inclination decreased. Trunk inclination, except at 28° downward, tended to parallel the changes in treadmill inclination. The most dramatic and consistent change for the hind limb joint displacement patterns was that maximum extension during stance increased as the treadmill inclination increased from 28° downward to 28° upward. In contrast to an earlier study by Prost & Sussman (1969), we could find no evidence that squirrel monkeys are best adapted to run on upward inclines of about 16°. The utilization of diagonal sequence gaits on the upward inclines supports previous suggestions that the preference for these gaits in primates is associated with an evolutionary increase in climbing behaviors.     

Biomechanical modeling and sensitivity analysis of bipedal running ability. I. Extant taxa

I used a simple mathematical model of the inverse dynamics of locomotion to estimate the minimum muscle masses required to maintain quasi-static equilibrium about the four main limb joints at mid-stance of fast running. Models of 10 extant taxa (a human, a kangaroo, two lizards, an alligator, and five birds) were analyzed in various bipedal poses to examine how anatomy, size, limb orientation, and other model parameters influence running ability. I examined how the muscle masses required for fast running compare to the muscle masses that are actually able to exert moments about the hip, knee, ankle, and toe joints, to see how support ability varies across the limb. I discuss the assumptions and limitations of the models, using sensitivity analysis to see how widely the results differed with feasible parameter input values. Even with a wide range of input values, the models validated the analysis procedure. Animals that are known to run bipedally were calculated as able to preserve quasi-static equilibrium about their hindlimb joints at mid-stance, whereas non-bipedal runners (iguanas and alligators) were recognized as having too little muscle mass to run quickly in bipedal poses. Thus, this modeling approach should be reliable for reconstructing running ability in extinct bipeds such as nonavian dinosaurs. The models also elucidated how key features are important for bipedal running capacity, such as limb orientation, muscle moment arms, muscle fascicle lengths, and body size. None of the animals modeled had extensor muscle masses acting about any one joint that were 7% or more of their body mass, which provides a reasonable limit for how much muscle mass is normally apportioned within a limb to act about a particular joint. The models consistently showed that a key biomechanical limit on running ability is the capacity of ankle extensors to generate sufficiently large joint moments. Additionally, the analysis reveals how large ratite birds remain excellent runners despite their larger size; they have apomorphically large extensor muscles with relatively high effective mechanical advantage. Finally, I reconstructed the evolution of running ability in the clade Reptilia, showing that the ancestors of extant birds likely were quite capable runners, even though they had already reduced key hip extensors such as M. caudofemoralis longus.     

Phylogenetic comparisons of pedestrian locomotion costs: confirmations and new insights

The energetic costs for animals to locomote on land influence many aspects of their ecology. Size accounts for much of the among‐species variation in terrestrial transport costs, but species of similar body size can still exhibit severalfold differences in energy expenditure. We compiled measurements of the (mass‐specific) minimum cost of pedestrian transport (COT_min, mL/kg/m) for 201 species – by far the largest sample to date – and used phylogenetically informed comparative analyses to investigate possible eco‐evolutionary differences in COT_min between various groupings of those species. We investigated number of legs, ectothermy and endothermy, waddling, and nocturnality specifically in lizards. Thus, our study primarily revisited previous theories about variations in COT_min between species, testing them with much more robust analyses. Having accounted for mass, while residual COT_min did not differ between bipedal and other species, specifically waddling bipeds were found to have relatively high COT_min. Furthermore, nocturnal lizards have relatively low COT_min although temperature does not appear to affect COT_min in ectotherms. Previous studies examining across‐species variation in COT_min from a biomechanical perspective show that the differences between waddling birds and nonwaddling species, and between nocturnal lizards and other ecotherms, are likely to be attributable to differences in ground reaction forces, posture, and effective limb length.     

How to climb a tree: lizards accelerate faster,but pause more,when escaping on vertical surfaces

Many species of lizards effectively traverse both two and three‐dimensional habitats. However, few studies have examined maximum locomotor performance on different inclines. Do maximum acceleration and velocity differ on a level and inclined surface? Do lizards pause more on an inclined surface? To address these questions, Sceloporus woodi lizards (N = 12) were run in the laboratory on a level trackway and a vertical tree trunk. This species is known to frequently utilize both vertical and horizontal aspects of its habitat. Average maximum acceleration on the vertical surface exceeded that on the level surface, although average maximum velocity exhibited the opposite pattern. The average number of pauses during level locomotion was lower compared to vertical locomotion. In addition, the average location of the first pause on the level surface was 0.51 m, which is farther than the average for vertical locomotion where the first pause was at 0.35 m. The combination of performance and pause data suggests that the relative lack of pausing during level locomotion allows individuals to reach higher maximum velocities on level surfaces because they accelerate over greater distances. The increased pausing when moving vertically could be a result of high energetic demands of vertical locomotion, or greater microhabitat complexity as a result of branching and/or refuges. The faster acceleration exhibited during vertical locomotion by S. woodi likely offsets the frequent pauses. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 83–90.     

The rate of metabolism in marine animals: environmental constraints, ecological demands and energetic opportunities

The rates of metabolism in animals vary tremendously throughout the biosphere. The origins of this variation are a matter of active debate with some scientists highlighting the importance of anatomical or environmental constraints, while others emphasize the diversity of ecological roles that organisms play and the associated energy demands. Here, we analyse metabolic rates in diverse marine taxa, with special emphasis on patterns of metabolic rate across a depth gradient, in an effort to understand the extent and underlying causes of variation. The conclusion from this analysis is that low rates of metabolism, in the deep sea and elsewhere, do not result from resource (e.g. food or oxygen) limitation or from temperature or pressure constraint. While metabolic rates do decline strongly with depth in several important animal groups, for others metabolism in abyssal species proceeds as fast as in ecologically similar shallow-water species at equivalent temperatures. Rather, high metabolic demand follows strong selection for locomotory capacity among visual predators inhabiting well-lit oceanic waters. Relaxation of this selection where visual predation is limited provides an opportunity for reduced energy expenditure. Large-scale metabolic variation in the ocean results from interspecific differences in ecological energy demand.     

Oxygen uptake kinetics in treadmill running and cycle ergometry: a comparison.

The purpose of the present study was to comprehensively examine oxygen consumption (VO(2)) kinetics during running and cycling through mathematical modeling of the breath-by-breath gas exchange responses to moderate and heavy exercise. After determination of the lactate threshold (LT) and maximal oxygen consumption (VO(2 max)) in both cycling and running exercise, seven subjects (age 26.6 +/- 5.1 yr) completed a series of "square-wave" rest-to-exercise transitions at running speeds and cycling power outputs that corresponded to 80% LT and 25, 50, and 75%Delta (Delta being the difference between LT and VO(2 max)). VO(2) responses were fit with either a two- (LT) exponential model. The parameters of the VO(2) kinetic response were similar between exercise modes, except for the VO(2) slow component, which was significantly (P < 0.05) greater for cycling than for running at 50 and 75%Delta (334 +/- 183 and 430 +/- 159 ml/min vs. 205 +/- 84 and 302 +/- 154 ml/min, respectively). We speculate that the differences between the modes are related to the higher intramuscular tension development in heavy cycle exercise and the higher eccentric exercise component in running. This may cause a relatively greater recruitment of the less efficient type II muscle fibers in cycling.     

Rainbow trout Oncorhynchus mykiss consume less energy when swimming near obstructions

The effect of obstructions in steady flow on swimming by rainbow trout Oncorhynchus mykiss was examined in a respirometry swim tunnel to test the prediction that fish interacting with obstructions require less energy to hold station. When an obstruction was present, O. mykiss altered the kinematics of swimming and the rate of oxygen consumption was significantly reduced. The fish employed both entrainment and Kármán gait swimming strategies, permitting greater locomotor efficiency.     

Aerobic and anaerobic determinants of repeated sprint ability in team sports athletes

The aim of this study was to examine in team sports athletes the relationship between repeated sprint ability (RSA) indices and both aerobic and anaerobic fitness components. Sixteen team-sport players were included (age, 23.4 ± 2.3 years; weight, 71.2 ± 8.3 kg; height, 178 ± 7 cm; body mass index, 22.4 ± 2 kg · m⁻²; estimated VO₂max, 54.16 ± 3.5 mL · kg⁻¹ · min⁻¹). Subjects were licensed in various team sports: soccer (n = 8), basketball (n = 5), and handball (n = 3). They performed 4 tests: the 20 m multi-stage shuttle run test (MSRT), the 30-s Wingate test (WingT), the Maximal Anaerobic Shuttle Running Test (MASRT), and the RSA test (10 repetitions of 30 m shuttle sprints (15 + 15 m with 180° change of direction) with 30 s passive recovery in between). Pearson''s product moment of correlation among the different physical tests was performed. No significant correlations were found between any RSA test indices and WingT. However, negative correlations were found between MASRT and RSA total sprint time (TT) and fatigue index (FI) (r = -0.53, p < 0.05 and r = -0.65, p < 0.01, respectively). No significant relationship between VO₂max and RSA peak sprint time (PT) and total sprint time (TT) was found. Nevertheless, VO₂max was significantly correlated with the RSA FI (r = -0.57, p < 0.05). In conclusion, aerobic fitness is an important factor influencing the ability to resist fatigue during RSA exercise. Our results highlighted the usefulness of MASRT, in contrast to WingT, as a specific anaerobic testing procedure to identify the anaerobic energy system contribution during RSA.     

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.