Why do we transition from walking to running? Energy cost and lower leg muscle activity before and after gait transition under body weight support.

BACKGROUND: Minimization of the energetic cost of transport (CoT) has been suggested for the walk-run transition in human locomotion. More recent literature argues that lower leg muscle activities are the potential triggers of the walk-run transition. We examined both metabolic and muscular aspects for explaining walk-run transition under body weight support (BWS; supported 30% of body weight) and normal walking (NW), because the BWS can reduce both leg muscle activity and metabolic rate. METHODS: Thirteen healthy young males participated in this study. The energetically optimal transition speed (EOTS) was determined as the intersection between linear CoT and speed relationship in running and quadratic CoT-speed relationship in walking under BWS and NW conditions. Preferred transition speed (PTS) was determined during constant acceleration protocol (velocity ramp protocol at 0.00463 m·s-2 = 1 km·h-1 per min) starting from 1.11 m·s-1. Muscle activities and mean power frequency (MPF) were measured using electromyography of the primary ankle dorsiflexor (tibialis anterior; TA) and synergetic plantar flexors (calf muscles including soleus) before and after the walk-run transition. RESULTS: The EOTS was significantly faster than the PTS under both conditions, and both were faster under BWS than in NW. In both conditions, MPF decreased after the walk-run transition in the dorsiflexor and the combined plantar flexor activities, especially the soleus. DISCUSSION: The walk-run transition is not triggered solely by the minimization of whole-body energy expenditure. Walk-run transition is associated with reduced TA and soleus activities with evidence of greater slow twitch fiber recruitment, perhaps to avoid early onset of localized muscle fatigue.

Sex differences in respiratory and circulatory cost during hypoxic walking: potential impact on oxygen saturation.

Energy expenditure (EE) during treadmill walking under normal conditions (normobaric normoxia, 21% O2) and moderate hypoxia (13% O2) was measured. Ten healthy young men and ten healthy young women walked on a level (0°) gradient a range of speeds (0.67-1.67 m s-1). During walking, there were no significant differences in reductions in arterial oxygen saturation (SpO2) between the sexes. The hypoxia-induced increase in EE, heart rate (HR [bpm]) and ventilation ([Formula: see text] [L min-1]) were calculated. Using a multivariate model that combined EE, [Formula: see text], and HR to predict ΔSpO2 (hypoxia-induced reduction), a very strong fit model both for men (r2 = 0.900, P < 0.001) and for women was obtained (r2 = 0.957, P < 0.001). The contributions of EE, VE, and HR to ΔSpO2 were markedly different between men and women. [Formula: see text] and EE had a stronger effect on ΔSpO2 in women ([Formula: see text]: 4.1% in women vs. 1.7% in men; EE: 28.1% in women vs. 15.8% in men), while HR had a greater effect in men (82.5% in men and 67.9% in women). These findings suggested that high-altitude adaptation in response to hypoxemia has different underlying mechanisms between men and women. These results can help to explain how to adapt high-altitude for men and women, respectively.

Combined effects of exposure to hypoxia and cool on walking economy and muscle oxygenation profiles at tibialis anterior.

We investigated combined effects of ambient temperature (23°C or 13°C) and fraction of inspired oxygen (21%O2 or 13%O2) on energy cost of walking (Cw: J·kg-1·km-1) and economical speed (ES). Eighteen healthy young adults (11 males, seven females) walked at seven speeds from 0.67 to 1.67 m s-1 (four min per stage). Environmental conditions were set; thermoneutral (N: 23°C) with normoxia (N: 21%O2) = NN; 23°C (N) with hypoxia (H: 13%O2) = NH; cool (C: 13°C) with 21%O2 (N) = CN, and 13°C (C) with 13%O2 (H) = CH. Muscle deoxygenation (HHb) and tissue O2 saturation (StO2) were measured at tibialis anterior. We found a significantly slower ES in NH (1.289 ± 0.091 m s-1) and CH (1.275 ± 0.099 m s-1) than in NN (1.334 ± 0.112 m s-1) and CN (1.332 ± 0.104 m s-1). Changes in HHb and StO2 were related to the ES. These results suggested that the combined effects (exposure to hypoxia and cool) is nearly equal to exposure to hypoxia and cool individually. Specifically, acute moderate hypoxia slowed the ES by approx. 4%, but acute cool environment did not affect the ES. Further, HHb and StO2 may partly account for an individual ES.

Impact of ambient temperature on energy cost and economical speed during level walking in healthy young males.

We measured oxygen consumption and carbon dioxide output during walking [per unit distance (Cw) values] for 14 healthy young human males at seven speeds from 0.67 to 1.67 m s-1 (4 min per stage) in thermoneutral (23°C), cool (13°C), and hot (33°C) environments. The Cw at faster gait speeds in the 33°C trial was slightly higher compared to those in the 23°C and 13°C trials. We found the speed at which the young males walked had a significant effect on the Cw values (P<0.05), but the different environmental temperatures showed no significant effect (P>0.05). Economical speed (ES) which can minimize the Cw in each individual was calculated from a U-shaped relationship. We found a significantly slower ES at 33°C [1.265 (0.060) m s-1 mean (s.d.)] compared to 23°C [1.349 (0.077) m s-1] and 13°C [1.356 (0.078) m s-1, P<0.05, respectively] with no differences between 23°C and 13°C (P>0.05). Heart rate and mean skin temperature responses in the 33°C condition increased throughout the walking trial compared to 23°C and 13°C (all P<0.05). These results suggest that an acutely hot environment slowed the ES by ∼7%, but an acutely cool environment did not affect the Cw and ES.

Energy cost and lower leg muscle activities during erect bipedal locomotion under hyperoxia.

BACKGROUND: Energy cost of transport per unit distance (CoT) against speed shows U-shaped fashion in walking and linear fashion in running, indicating that there exists a specific walking speed minimizing the CoT, being defined as economical speed (ES). Another specific gait speed is the intersection speed between both fashions, being called energetically optimal transition speed (EOTS). We measured the ES, EOTS, and muscle activities during walking and running at the EOTS under hyperoxia (40% fraction of inspired oxygen) on the level and uphill gradients (+ 5%). METHODS: Oxygen consumption [Formula: see text] and carbon dioxide output [Formula: see text] were measured to calculate the CoT values at eight walking speeds (2.4-7.3 km h-1) and four running speeds (7.3-9.4 km h- 1) in 17 young males. Electromyography was recorded from gastrocnemius medialis, gastrocnemius lateralis (GL), and tibialis anterior (TA) to evaluate muscle activities. Mean power frequency (MPF) was obtained to compare motor unit recruitment patterns between walking and running. RESULTS: [Formula: see text], [Formula: see text], and CoT values were lower under hyperoxia than normoxia at faster walking speeds and any running speeds. A faster ES on the uphill gradient and slower EOTS on both gradients were observed under hyperoxia than normoxia. GL and TA activities became lower when switching from walking to running at the EOTS under both FiO2 conditions on both gradients, so did the MPF in the TA. CONCLUSIONS: ES and EOTS were influenced by reduced metabolic demands induced by hyperoxia. GL and TA activities in association with a lower shift of motor unit recruitment patterns in the TA would be related to the gait selection when walking or running at the EOTS. TRIAL REGISTRATION: UMIN000017690 ( R000020501 ). Registered May 26, 2015, before the first trial.

Walking economy at simulated high altitude in human healthy young male lowlanders.

We measured oxygen consumption during walking per unit distance (Cw) values for 12 human healthy young males at six speeds from 0.667 to 1.639 m s-1 (four min per stage) on a level gradient under normobaric normoxia, moderate hypoxia (15% O2), and severe hypoxia (11% O2). Muscle deoxygenation (HHb) was measured at the vastus lateralis muscle using near-infrared spectroscopy. Economical speed which can minimize the Cw in each individual was calculated from a U-shaped relationship. We found a significantly slower economical speed (ES) under severe hypoxia [1.237 (0.056) m s-1; mean (s.d.)] compared to normoxia [1.334 (0.070) m s-1] and moderate hypoxia [1.314 (0.070) m s-1, P<0.05 respectively] with no differences between normoxia and moderate hypoxia (P>0.05). HHb gradually increased with increasing speed under severe hypoxia, while it did not increase under normoxia and moderate hypoxia. Changes in HHb between standing baseline and the final minute at faster gait speeds were significantly related to individual ES (r=0.393 at 1.250 m s-1, r=0.376 at 1.444 m s-1, and r=0.409 at 1.639 m s-1, P<0.05, respectively). These results suggested that acute severe hypoxia slowed ES by ∼8%, but moderate hypoxia left ES unchanged.

Economical Speed and Energetically Optimal Transition Speed Evaluated by Gross and Net Oxygen Cost of Transport at Different Gradients.

The oxygen cost of transport per unit distance (CoT; mL·kg(-1)·km(-1)) shows a U-shaped curve as a function of walking speed (v), which includes a particular walking speed minimizing the CoT, so called economical speed (ES). The CoT-v relationship in running is approximately linear. These distinctive walking and running CoT-v relationships give an intersection between U-shaped and linear CoT relationships, termed the energetically optimal transition speed (EOTS). This study investigated the effects of subtracting the standing oxygen cost for calculating the CoT and its relevant effects on the ES and EOTS at the level and gradient slopes (±5%) in eleven male trained athletes. The percent effects of subtracting the standing oxygen cost (4.8 ± 0.4 mL·kg(-1)·min(-1)) on the CoT were significantly greater as the walking speed was slower, but it was not significant at faster running speeds over 9.4 km·h(-1). The percent effect was significantly dependent on the gradient (downhill > level > uphill, P < 0.001). The net ES (level 4.09 ± 0.31, uphill 4.22 ± 0.37, and downhill 4.16 ± 0.44 km·h(-1)) was approximately 20% slower than the gross ES (level 5.15 ± 0.18, uphill 5.27 ± 0.20, and downhill 5.37 ± 0.22 km·h(-1), P < 0.001). Both net and gross ES were not significantly dependent on the gradient. In contrast, the gross EOTS was slower than the net EOTS at the level (7.49 ± 0.32 vs. 7.63 ± 0.36 km·h(-1), P = 0.003) and downhill gradients (7.78 ± 0.33 vs. 8.01 ± 0.41 km·h(-1), P < 0.001), but not at the uphill gradient (7.55 ± 0.37 vs. 7.63 ± 0.51 km·h(-1), P = 0.080). Note that those percent differences were less than 2.9%. Given these results, a subtraction of the standing oxygen cost should be carefully considered depending on the purpose of each study.

Effects of load and gradient on energy cost of running.

This study quantified the interaction of electromyography (EMG) obtained from the vastus lateralis and metabolic energy cost of running (C(r); mL·[mass+load](-1)·meter(-1)), an index of running economy, during submaximal treadmill running. Experiments were conducted with and without load on the back on a motor-driven treadmill on the downhill, level and uphill slopes. The obtained EMG was full-wave rectified and integrated (iEMG). The iEMG was divided into eccentric (ECC) and concentric (CON) phases with a foot sensor and a knee-joint goniometer. The ratio of ECC to CON (ECC/CON ratio) was regarded as the muscle elastic capacity during running on each slope. The C(r) was determined as the ratio of the 2-min steady-state VO(2) to the running speed. We found a significant decrease in the C(r) when carrying the load at all slopes. The ECC/CON ratio was significantly higher in the load condition at the downhill and level slopes, but not at the uphill slope. A significant gradient difference was observed in the C(r) (downuphill). Thus, an alteration of Cr by the gradient and load was almost consistent with that of the ECC/CON ratio. The ECC/CON ratio, but not the rotative torque (T) functioning around the center of body mass, significantly correlated with C(r) (r=-0.41, p<0.05). These results indicated that the ECC/CON ratio, rather than T, contributed to one of the energy-saving mechanisms during running with load.

Changes in EMG characteristics and metabolic energy cost during 90-min prolonged running.

This study quantified the interaction of integrated electromyography (iEMG) obtained from the vastus lateralis and the metabolic energy cost of running (Cr), an index to assess running economy, during 90-min prolonged running. The iEMG during running was divided into eccentric (ECC) and concentric (CON) phases using a force platform and a knee-joint goniometer. The ratio of ECC to CON (ECC/CON ratio) significantly decreased during 90-min prolonged running in novice distance runners, which would be explained by an increase in muscle activity during the CON phase of running. The average Cr value significantly increased during 90-min prolonged running. The individual's Cr values significantly correlated with the ECC/CON ratio (r=-0.702, P<0.05). These results suggest that changes in the ECC/CON ratio and Cr value during prolonged running are associated.

Assessment of short-distance breaststroke swimming performance with critical velocity.

For high-velocity running or swimming, the relationship between velocity (v) and its sustainable duration (t) can be described by a hyperbolic relationship: (v - Vcrit)·t = D', where Vcrit is termed critical velocity, and D' is defined as a curvature constant of the hyperbolic curve. The purposes of this study were to examine whether the Vcrit could be applied to evaluate short-distance breaststroke swimming performance and to evaluate the relative contribution of D' in short-distance swimming performance. Eleven male swimmers performed a series of time trials corresponding to 75, 100, and 150-m in an indoor 50-m swimming pool. The observed records were calculated into average velocities of each event to determine Vcrit and D'. After the determination of Vcrit and D', all subjects performed 50-m time trial on another day. A maximal anaerobic power test using cycle ergometer was also performed in the laboratory. The average velocity of the 50-m time trial significantly correlated with the obtained Vcrit, but not with D'. D' was significantly correlated with the residual error, calculated from the regression analysis for the relationship between Vcrit and the average velocities of 50-m time trial. A cluster analysis showed that most of the subjects were classified as Vcrit dependency when performing 50-m time trial. Those results indicated that Vcrit could be applied to evaluate short-distance swimming performance, and it determined around 80% of the short-distance breaststroke swimming performance. Key PointsFor high-velocity running or swimming, the relationship between velocity (v) and its sustainable duration (t) can be described by a hyperbolic relationship: (v - Vcrit)·t = D', where Vcrit is termed critical velocity, and D' is defined as a curvature constant of the hyperbolic curve. The D' contributed only around 20% of the breaststroke swimming performance even in a short-distance event.Critical velocity determined around 80% of 50-m breaststroke swimming performance, and it could be a useful tool for evaluating short-distance swimming performance.Most of the swimmers showed characteristics for critical velocity dependent physical fitness even in short-distance swimming event.