Search

This comment addresses the incomplete presentation and incorrect conclusion offered in the recent manuscript of Beck et al . ( R. Soc. Open Sci. 9 , 211799 (doi:10.1098/rsos.211799)). The manuscript introduces biomechanical and performance data on the fastest-ever, bilateral amputee 400 m runner. Using an advantage standard of not faster than the fastest non-amputee runner ever (i.e. performance superior to that of the intact-limb world record-holder), the Beck et al . manuscript concludes that sprint running performance on bilateral, lower-limb prostheses is not unequivocally advantageous compared to the biological limb condition. The manuscript acknowledges the long-standing support of the authors for the numerous eligibility applications of the bilateral-amputee athlete. However, it does not acknowledge that the athlete's anatomically disproportionate prosthetic limb lengths (+15 cm versus the World Para Athletics maximum) are ineligible in both Olympic and Paralympic track competition due to their performance-enhancing properties. Also not acknowledged are the slower sprint performances of the bilateral-amputee athlete on limbs of shorter length that directly refute their manuscript's primary conclusion. Our contribution here provides essential background information and data not included in the Beck et al. manuscript that make the correct empirical conclusion clear: artificially long legs artificially enhance long sprint running performance., (© 2022 The Authors.)

The durations that muscular force and power outputs can be sustained until failure fall predictably on an exponential decline between an individual's 3-s burst maximum to the maximum performance they can sustain aerobically. The exponential time constants describing these rates of performance loss are similar across individuals, suggesting that a common metabolically based mechanism governs muscle fatigue; however, these conclusions come from studies mainly on men. To test whether the same physiological understanding can be applied to women, we compared the performance-duration relationships and neuromuscular activity between seven men [23.3 ± 1.9 (SD) yr] and seven women (21.7 ± 1.8 yr) from multiple exhaustive bouts of cycle ergometry. Each subject performed trials to obtain the peak 3-s power output (P max ), the mechanical power at the aerobic maximum (P aer ), and 11-14 constant-load bouts eliciting failure between 3 and 300 s. Collectively, men and women performed 180 exhaustive bouts spanning an ~6-fold range of power outputs (118-1116 W) and an ~35-fold range of trial durations (8-283 s). Men generated 66% greater P max (956 ± 109 W vs. 632 ± 74 W) and 68% greater P aer (310 ± 47 W vs. 212 ± 15 W) than women. However, the metabolically based time constants describing the time course of performance loss were similar between men (0.020 ± 0.003/s) and women (0.021 ± 0.003/s). Additionally, the fatigue-induced increases in neuromuscular activity did not differ between the sexes when compared relative to the pedal forces at P aer These data suggest that muscle fatigue during short-duration dynamic exercise has a common metabolically based mechanism determined by the extent that ATP is resynthesized by anaerobic metabolism., New & Noteworthy: Although men and women differed considerably in their absolute cycling performances, there was no sex difference in the metabolically based exponential time constant that described the performance-duration relationship. Similarly, the fatigue-induced increases in neuromuscular activity were not different between the sexes when compared from a metabolic perspective. These data suggest that men and women have similar rate-limiting mechanisms for short-duration dynamic exercise that are determined by the extent the exercise is supported by anaerobic metabolism., (Copyright © 2017 the American Physiological Society.)

Investigations focused on the gait and physiological limits of human speed have been on-going for more than a century. However, due to measurement limitation a kinetic understanding of the foot-ground collision and how these dynamics differ between individuals to confer speed and limit gait has only recently begun to come forth. Therefore, we designed and tested an instrumented high-speed force treadmill to measure the forces occurring at the limits of human performance. The treadmill was designed to maximize flexural stiffness and natural frequency by using a honeycomb sandwich panel as the bed surface and a flexible drive shaft between the drive roller and servo motor to reduce the mass of the supported elements which contribute to the system's response frequency. The functional performance of the force treadmill met or exceeded the measurement criteria established for ideal force plates: high natural frequency (z-axis = 113 Hz), low crosstalk between components of the force (Fx/Fz = 0.0020[SD = 0.0010]; Fy/Fz = 0.0016[SD = 0.0003]), a linear response (R(2) > 0.999) for loading with known weights (range: 44-3857 N), and an accuracy of 2.5[SD = 1.7] mm and 2.8[SD = 1.5] mm in the x and y-axes, respectively, for the point of force application. In dynamic testing at running speeds up to 10 m s(-1), the measured durations and magnitudes of force application were similar between the treadmill and over-ground running using a force platform. This design provides a precise instrumented treadmill capable of recording multi-axis ground reaction forces applied during the foot ground contacts of the fastest men and animals known to science., (Copyright © 2015 IPEM. Published by Elsevier Ltd. All rights reserved.)

We investigated the influence of altered muscle duty cycle on the performance decrements and neuromuscular responses occurring during constant-load, fatiguing bouts of knee extension exercise. We experimentally altered the durations of the muscularly inactive portion of the limb movement cycle and hypothesized that greater relative durations of inactivity within the same movement task would 1) reduce the rates and extent of muscle performance loss and 2) increase the forces necessary to trigger muscle fatigue. In each condition (duty cycle = 0.6 and 0.3), male subjects [age = 25.9 ± 2.0 yr (SE); mass = 85.4 ± 2.6 kg], completed 9-11 exhaustive bouts of two-legged knee extension exercise, at force outputs that elicited failure between 4 and 290 s. The novel duty cycle manipulation produced two primary results; first, we observed twofold differences in both the extent of muscle performance lost (DC0.6 = 761 ± 35 N vs. DC0.3 = 366 ± 49 N) and the time course of performance loss. For example, exhaustive trials at the midpoint of these force ranges differed in duration by more than 30 s (t0.6 = 36 ± 2.6 vs. t0.3 = 67 ± 4.3 s). Second, both the minimum forces necessary to exceed the peak aerobic capacity and initiate a reliance on anaerobic metabolism, and the forces necessary to elicit compensatory increases in electromyogram activity were 300% greater in the lower vs. higher duty cycle condition. These results indicate that the fatigue-induced compensatory behavior to recruit additional motor units is triggered by a reliance on anaerobic metabolism for ATP resynthesis and is independent of the absolute level or fraction of the maximum force produced by the muscle., (Copyright © 2015 the American Physiological Society.)

Prevailing physiological paradigms explain both sprint and endurance exercise performance in terms of the availability of metabolic energy. However, for all-out efforts of 60 s or less, the prevailing view is no longer viable. Contemporary evidence indicates that sprinting performance is determined by musculoskeletal force application, with a duration dependency explained by the intrinsically rapid rates at which skeletal muscle fatigues in vivo.

Running speed is limited by a mechanical interaction between the stance and swing phases of the stride. Here, we tested whether stance phase limitations are imposed by ground force maximums or foot-ground contact time minimums. We selected one-legged hopping and backward running as experimental contrasts to forward running and had seven athletic subjects complete progressive discontinuous treadmill tests to failure to determine their top speeds in each of the three gaits. Vertical ground reaction forces [in body weights (W(b))] and periods of ground force application (T(c); s) were measured using a custom, high-speed force treadmill. At top speed, we found that both the stance-averaged (F(avg)) and peak (F(peak)) vertical forces applied to the treadmill surface during one-legged hopping exceeded those applied during forward running by more than one-half of the body's weight (F(avg) = 2.71 +/- 0.15 vs. 2.08 +/- 0.07 W(b); F(peak) = 4.20 +/- 0.24 vs. 3.62 +/- 0.24 W(b); means +/- SE) and that hopping periods of force application were significantly longer (T(c) = 0.160 +/- 0.006 vs. 0.108 +/- 0.004 s). Next, we found that the periods of ground force application at top backward and forward running speeds were nearly identical, agreeing to within an average of 0.006 s (T(c) = 0.116 +/- 0.004 vs. 0.110 +/- 0.005 s). We conclude that the stance phase limit to running speed is imposed not by the maximum forces that the limbs can apply to the ground but rather by the minimum time needed to apply the large, mass-specific forces necessary.

The University of Wyomings Biomechanics Laboratory is examining the link between the energetics and mechanics of both human and animal locomotion. The group is currently investigating how there lative duration of muscular activity (i.e. duty factor) influences the time course of performance loss resulting from the impairment in muscle force production that occurs as the duration of an all-out or text ends from seconds to minutes. In order to conduct one of their experimental protocols, this group requires human subjects to precisely and repeatedly match prescribed contractile and relaxation periods during knee-extension exercise. To provide subjects with visual feedback with respect to their adherence to the assigned contractile durations and limb velocities, a custom instrumentation array panel was developed consisting of two arrays of high-visibility light emitting diodes. One array provides the experimental subject with the selected, or target, rate of position change while the complementary array provides an indication of the actual rate of displacement achieved by the subject.

The recent competitive successes of a bilateral, transtibial amputee sprint runner who races with modern running prostheses has triggered an international controversy regarding the relative function provided by his artificial limbs. Here, we conducted three tests of functional similarity between this amputee sprinter and competitive male runners with intact limbs: the metabolic cost of running, sprinting endurance, and running mechanics. Metabolic and mechanical data, respectively, were acquired via indirect calorimetry and ground reaction force measurements during constant-speed, level treadmill running. First, we found that the mean gross metabolic cost of transport of our amputee sprint subject (174.9 ml O(2)*kg(-1)*km(-1); speeds: 2.5-4.1 m/s) was only 3.8% lower than mean values for intact-limb elite distance runners and 6.7% lower than for subelite distance runners but 17% lower than for intact-limb 400-m specialists [210.6 (SD 13.2) ml O(2)*kg(-1)*km(-1)]. Second, the speeds that our amputee sprinter maintained for six all-out, constant-speed trials to failure (speeds: 6.6-10.8 m/s; durations: 2-90 s) were within 2.2 (SD 0.6)% of those predicted for intact-limb sprinters. Third, at sprinting speeds of 8.0, 9.0, and 10.0 m/s, our amputee subject had longer foot-ground contact times [+14.7 (SD 4.2)%], shorter aerial [-26.4 (SD 9.9)%] and swing times [-15.2 (SD 6.9)%], and lower stance-averaged vertical forces [-19.3 (SD 3.1)%] than intact-limb sprinters [top speeds = 10.8 vs. 10.8 (SD 0.6) m/s]. We conclude that running on modern, lower-limb sprinting prostheses appears to be physiologically similar but mechanically different from running with intact limbs.

Based on aerodynamic considerations, the energy use-flight speed relationship of all airborne animals and aircraft should be U-shaped. However, measures of the metabolic rate-flight speed relationship in birds have been available since Tucker's pioneering experiments with budgerigars nearly forty years ago, but this classic work remains the only study to have found a clearly U-shaped metabolic power curve. The available data suggests that the energetic requirements for flight within this species are unique, yet the metabolic power curve of the budgerigar is widely considered representative of birds in general. Given these conflicting results and the observation that the budgerigar's mass is less than 50% of the next smallest species to have been studied, we asked whether large and small birds have metabolic power curves of different shapes. To address this question we measured the rates of oxygen uptake and wingbeat kinematics in budgerigars and cockatiels flying within a variable-speed wind tunnel. These species are close phylogenetic relatives, have similar flight styles, wingbeat kinematics, and are geometrically similar but have body masses that differ by a factor of two. In contrast to our expectations, we found the metabolic rate-flight speed relationship of both species to be acutely U-shaped. We also found that neither budgerigars nor cockatiels used their normal intermittent flight style while wearing a respirometric mask. We conclude that species size differences alone do not explain the previously unique metabolic power curve of the budgerigar; however, due to the absence of comparable data we cannot evaluate whether the mask-related kinematic response we document influences the metabolic rate-flight speed relationship of these parrots, or whether the energetics of flight differ between this and other avian clades.

For both different individuals and modes of locomotion, the external forces determining all-out sprinting performances fall predictably with effort duration from the burst maximums attained for 3 s to those that can be supported aerobically as trial durations extend to roughly 300 s. The common time course of this relationship suggests a metabolic basis for the decrements in the force applied to the environment. However, the mechanical and neuromuscular responses to impaired force production (i.e., muscle fatigue) are generally considered in relation to fractions of the maximum force available, or the maximum voluntary contraction (MVC). We hypothesized that these duration-dependent decrements in external force application result from a reliance on anaerobic metabolism for force production rather than the absolute force produced. We tested this idea by examining neuromuscular activity during two modes of sprint cycling with similar external force requirements but differing aerobic and anaerobic contributions to force production: one- and two-legged cycling. In agreement with previous studies, we found greater peak per leg aerobic metabolic rates [59% (+/-6 SD)] and pedal forces at VO2 peak [30% (+/-9)] during one- vs. two-legged cycling. We also determined downstroke pedal forces and neuromuscular activity by surface electromyography during 15 to 19 all-out constant load sprints lasting from 12 to 400 s for both modes of cycling. In support of our hypothesis, we found that the greater reliance on anaerobic metabolism for force production induced compensatory muscle recruitment at lower pedal forces during two- vs. one-legged sprint cycling. We conclude that impaired muscle force production and compensatory neuromuscular activity during sprinting are triggered by a reliance on anaerobic metabolism for force production.

We hypothesized that the maximum mechanical power outputs that can be maintained during all-out sprint cycling efforts lasting from a few seconds to several minutes can be accurately estimated from a single exponential time constant (k(cycle)) and two measurements on individual cyclists: the peak 3-s power output (P(mech max)) and the maximum mechanical power output that can be supported aerobically (P(aer)). Tests were conducted on seven subjects, four males and three females, on a stationary cycle ergometer at a pedal frequency of 100 rpm. Peak mechanical power output (P(mech max)) was the highest mean power output attained during a 3-s burst; the maximum power output supported aerobically (P(aer)) was determined from rates of oxygen uptake measured during a progressive, discontinuous cycling test to failure. Individual power output-duration relationships were determined from 13 to 16 all-out constant load sprints lasting from 5 to 350 s. In accordance with the above hypothesis, the power outputs measured during all-out sprinting efforts were estimated to within an average of 34 W or 6.6% from P(mech max), P(aer), and a single exponential constant (k(cycle) = 0.026 s(-1)) across a sixfold range of power outputs and a 70-fold range of sprint trial durations (R2 = 0.96 vs. identity, n = 105; range: 180 to 1,136 W). Duration-dependent decrements in sprint cycling power outputs were two times greater than those previously identified for sprint running speed (k(run) = 0.013 s(-1)). When related to the respective times of pedal and ground force application rather than total sprint time, decrements in sprint cycling and running performance followed the same time course (k = 0.054 s(-1)). We conclude that the duration-dependent decrements in sprinting performance are set by the fractional duration of the relevant muscular contractions.

We hypothesized that the anaerobic power and aerobic power outputs during all-out runs of any common duration between 10 and 150 s would be proportional to the maximum anaerobic (E(an-max)) and aerobic powers (E(aer-max)) available to the individual runner. Seventeen runners who differed in E(an-max) and E(aer-max) (5 sprinters, 5 middle-distance runners, and 7 long distance runners) were tested during treadmill running on a 4.6 degrees incline. E(an-max) was estimated from the fastest treadmill speed subjects could attain for eight steps. E(aer-max) was determined from a progressive, discontinuous, treadmill test to failure. Oxygen deficits and rates of uptake were measured to assess the respective anaerobic and aerobic power outputs during 11-16 all-out treadmill runs that elicited failure between 10 and 220 s. We found that, during all-out runs of any common duration, the relative anaerobic and aerobic powers utilized were largely the same for sprint, middle-distance, and long-distance subjects. The similar fractional utilization of the E(an-max) and E(aer-max) available during high-speed running 1) provides empirical values that modify and advance classic theory, 2) allows rates of anaerobic and aerobic energy release to be quantified from individual maxima and run durations, and 3) explains why the high-speed running performances of different event specialists can be accurately predicted (R(2) = 0.97; n = 254) from two direct measurements and the same exponential time constant.

A recently discovered locomotor behavior, wing-assisted incline running (WAIR), allows fully volant animals to 'run' up vertical obstacles. Such a task would appear to be especially formidable for bipeds, yet WAIR is used preferentially by ground-dwelling birds, specifically chukar partridge Alectoris chukar, to reach refugia. The basic locomotor mechanics that enable this behavior are not fully understood. For instance, are there functional differences at the level of the wing during WAIR and free flight, and do the hindlimbs actively participate in propulsion during WAIR? To investigate wing function during these activities we used accelerometry to compare the instantaneous whole-body acceleration during WAIR and ascending free flights at a similar climb angle. Throughout a substantial portion of the wingbeat cycle, chukars engaged in WAIR experienced an acceleration oriented towards the substrate, whereas during ascending free flights the acceleration of the center of mass was parallel to the direction of travel. We investigated whether the animals were using their hindlimbs for propulsion, rather than for some other function (e.g. to maintain balance), by measuring ground reaction forces (GRF) during bouts of WAIR. Estimates of the contribution of the hindlimbs towards the vertical external work done by the bird were 98 +/- 8% of the total at an incline of 60 degrees (the steepest angle that birds were able to negotiate without the use of their forelimbs). During vertical (90 degrees ) bouts of WAIR the hindlimb contribution was 37 +/- 5% of the total external work. Yet, the magnitude of the peak GRF at 90 degrees was 175% of the value generated during level walking, revealing that birds engaged in WAIR do generate sizeable hindlimb forces even during vertical ascents. These data support the hypothesis that forelimbs are enabling hindlimb function, and we argue that this represents a locomotor strategy which may have been used by proto-birds during the evolution of flight.

We hypothesized that all-out running speeds for efforts lasting from a few seconds to several minutes could be accurately predicted from two measurements: the maximum respective speeds supported by the anaerobic and aerobic powers of the runner. To evaluate our hypothesis, we recruited seven competitive runners of different event specialties and tested them during treadmill and overground running on level surfaces. The maximum speed supported by anaerobic power was determined from the fastest speed that subjects could attain for a burst of eight steps (approximately 3 s or less). The maximum speed supported by aerobic power, or the velocity at maximal oxygen uptake, was determined from a progressive, discontinuous treadmill test to failure. All-out running speeds for trials of 3-240 s were measured during 10-13 constant-speed treadmill runs to failure and 4 track runs at specified distances. Measured values of the maximum speeds supported by anaerobic and aerobic power, in conjunction with an exponential constant, allowed us to predict the speeds of all-out treadmill trials to within an average of 2.5% (R2 = 0.94; n = 84) and track trials to within 3.4% (R2 = 0.86; n = 28). An algorithm using this exponent and only two of the all-out treadmill runs to predict the remaining treadmill trials was nearly as accurate (average = 3.7%; R2 = 0.93; n = 77). We conclude that our technique 1) provides accurate predictions of high-speed running performance in trained runners and 2) offers a performance assessment alternative to existing tests of anaerobic power and capacity.

While useful in describing the efficiency of maneuvering flight, steady-state (i.e., fixed wing) models of maneuvering performance cannot provide insight to the efficacy of maneuvering, particularly during low-speed flapping flight. Contrasted with airplane-analogous gliding/high speed maneuvering, the aerodynamic and biomechanical mechanisms employed by birds at low flight speeds are violent, with rapidly alternating forces routinely being developed. The saltatory nature of this type of flight results in extreme linear and angular displacements of the bird's body; however, birds isolate their heads from these accelerations with cervical reflexes. Experiments with pigeons suggest this ability to isolate the visual and vestibular systems is critical to controlled flapping flight: birds wearing collars that prohibited the neck from isolating the head from the angular accelerations of induced rolls frequently exhibited (50% of flights) a loss of vestibular and/or visual horizon and were unable to maintain controlled flight.

We tested the importance of aerobic metabolism to human running speed directly by altering inspired oxygen concentrations and comparing the maximal speeds attained at different rates of oxygen uptake. Under both normoxic (20.93% O2) and hypoxic (13.00% O2) conditions, four fit adult men completed 15 all-out sprints lasting from 15 to 180 s as well as progressive, discontinuous treadmill tests to determine maximal oxygen uptake and the metabolic cost of steady-state running. Maximal aerobic power was lower by 30% (1.00 +/- 0.15 vs. 0.77 +/- 0.12 ml O2. kg-1. s-1) and sprinting rates of oxygen uptake by 12-25% under hypoxic vs. normoxic conditions while the metabolic cost of submaximal running was the same. Despite reductions in the aerobic energy available for sprinting under hypoxic conditions, our subjects were able to run just as fast for sprints of up to 60 s and nearly as fast for sprints of up to 120 s. This was possible because rates of anaerobic energy release, estimated from oxygen deficits, increased by as much as 18%, and thus compensated for the reductions in aerobic power. We conclude that maximal metabolic power outputs during sprinting are not limited by rates of anaerobic metabolism and that human speed is largely independent of aerobic power during all-out runs of 60 s or less.