What are the relations between mechanics, gait parameters, and energetics in terrestrial locomotion?

Are the different energy-conserving mechanics (i.e., pendulum and spring) used in different gaits reflected in differences in energetics and/or stride parameters? The analysis included published data from several species and new data from horses. When changing from pendulum to spring mechanics, there is a change in the slope of metabolic rate (MR) vs. speed in all species, in birds and quadrupeds there is no step increase, and in humans there are conflicting reports. At the trot-gallop transition, where quadrupeds are hypothesized to change from spring mechanics to some combination of spring and pendulum mechanics, there is a change in slope of MR vs. speed in horses but not in other species. Stride frequency (SF) is a logarithmic function of walking speed in all species, a linear function of trotting/running speed, and nearly independent of speed in galloping. In humans and horses there is a discontinuity in SF at the walk-trot (run) transition but not in birds. The slope of time of contact vs. speed does not change with mechanics in most species, but it does in humans. In horses and humans, there is a discontinuity at the walk-trot (run) transition and data for other species do not permit generalization. Duty factor (DF) in humans is greater than 0.5 in walking (pendulum mechanics) and less than 0.5 when running (spring mechanics). However, this is not true in many species that have DF>0.5 at the lowest speeds where they use spring mechanics. When trotting at low speeds, horses use forelimb DF>0.5 and hind limb DF<0.5. Thus, it is confusing to distinguish between walking and running by DF.

Joint work and power for both the forelimb and hindlimb during trotting in the horse.

The net work of the limbs during constant speed over level ground should be zero. However, the partitioning of negative and positive work between the fore- and hindlimbs of a quadruped is not likely to be equal because the forelimb produces a net braking force while the hindlimb produces a net propulsive force. It was hypothesized that the forelimb would do net negative work while the hindlimb did net positive work during trotting in the horse. Because vertical and horizontal impulses remain unchanged across speeds it was hypothesized that net work of both limbs would be independent of speed. Additionally because the major mass of limb musculature is located proximally, it was hypothesized that proximal joints would do more work than distal joints. Kinetic and kinematic analysis were combined using inverse dynamics to calculate work and power for each joint of horses trotting at between 2.5 and 5.0 m s(-1). Work done by the hindlimb was indeed positive (consistently 0.34 J kg(-1) across all speeds), but, contrary to our hypothesis, net work by the forelimb was essentially zero (but also independent of trotting speed). The zero net work of the forelimb may be the consequence of our not being able to account, experimentally, for the negative work done by the extrinsic muscles connecting the scapula and the thorax. The distal three joints of both limbs behaved elastically with a period of energy absorption followed by energy return. Proximal forelimb joints (elbow and shoulder) did no net work, because there was very little movement of the elbow and shoulder during the portion of stance when an extensor moment was greatest. Of the two proximal hindlimb joints, the hip did positive work during the stride, generating energy almost throughout stance. The knee did some work, but like the forelimb proximal joints, had little movement during the middle of stance when the flexion moment was the greatest, probably serving to allow the efficient transmission of energy from the hip musculature to the ground.

Ground reaction forces in horses trotting up an incline and on the level over a range of speeds.

Although the forces required to support the body mass are not elevated when moving up an incline, kinematic studies, in vivo tendon and bone studies and kinetic studies suggest there is a shift in forces from the fore- to the hindlimbs in quadrupeds. However, there are no whole-animal kinetic measurements of incline locomotion. Based on previous related research, we hypothesized that there would be a shift in forces to the hindlimb. The present study measured the force produced by the fore- and hindlimbs of horses while trotting over a range of speeds (2.5 to 5 m s(-1)) on both level and up an inclined (10%) surface. On the level, forelimb peak forces increased with trotting speed, but hindlimb peak force remained constant. On the incline, both fore- and hindlimb peak forces increased with speed, but the sum of the peak forces was lower than on the level. On the level, over the range of speeds tested, total force was consistently distributed between the limbs as 57% forelimb and 43% hindlimb, similar to the weight distribution of the horses during static weight tests. On the incline, the force distribution during locomotion shifted to 52% forelimb and 48% hindlimb. Time of contact and duty factor decreased with speed for both limbs. Time of contact was longer for the forelimb than the hindlimb, a finding not previously reported for quadrupeds. Time of contact of both limbs tended to be longer when traveling up the incline than on the level, but duty factor for both limbs was similar under both conditions. Duty factor decreased slightly with increased speed for the hindlimb on the level, and the corresponding small, predicted increase in peak vertical force could not be detected statistically.

DOMS-associated changes in ankle and knee joint dynamics during running.

PURPOSE: The purpose of this study was to determine whether leg mechanics change due to DOMS by examining ankle and knee joint kinematics and stiffness before and after a down hill run. METHODS: Sagittal plane kinematics were recorded with high-speed (120 Hz) video at a speed representing 75% of VO2peak of nine well-trained male runners before (RE1) and 48 h after (RE2) a 30-min downhill run. From the recorded video, 10-12 consecutive strides were digitized, and the following variables were calculated for each stride: ankle and knee range of motion (ROM), ankle and knee peak angular velocity, ankle and knee stiffness, and leg vertical stiffness. A repeated measures ANOVA was calculated for each variable (alpha = 0.05). RESULTS: Both knee and ankle ROM during stance decreased with DOMS, but otherwise there were few changes in ankle mechanics with DOMS. Knee stiffness tended to increase during the early portion of stance (from initial stance to maximum angular velocity of flexion) with DOMS, immediately followed by a decrease (to maximum knee flexion) in stiffness. Changes in knee stiffness caused vertical leg stiffness to increase for the initial portion of stance with DOMS. CONCLUSION: Knee mechanics changed such that the knee stiffness increased at initial stance, resulting in an increase in vertical leg stiffness. This change in knee stiffness possibly serves as a protective mechanism to prevent further damage or pain in the knee extensor musculature.

Moments and power generated by the horse (Equus caballus) hind limb during jumping.

The ability to jump over an obstacle depends upon the generation of work across the joints of the propelling limb(s). The total work generated by one hind limb of a horse and the contribution to the total work by four joints of the hind limb were determined for a jump. It was hypothesized that the hip and ankle joints would have extensor moments performing positive work, while the knee would have a flexor moment and perform negative work during the jump. Ground reaction forces and sagittal plane kinematics were simultaneously recorded during each jumping trial. Joint moment, power and work were determined for the metatarsophalangeal (MP), tarsal (ankle), tibiofemoral (knee) and coxofemoral (hip) joints. The hip, knee and ankle all flexed and then extended and the MP extended and then flexed during ground contact. Consistent with our hypothesis, large extensor moments were observed at the hip and ankle joints and large flexor moments at the knee and MP joints throughout ground contact of the hind limb. Peak moments tended to occur earlier in stance in the proximal joints but peak power generation of the hind limb joints occurred at similar times except for the MP joint, with the hip and ankle peaking first followed by the MP joint. During the first portion of ground contact (approximately 40%), the net result of the joint powers was the absorption of power. During the remainder of the contact period, the hind limb generated power. This pattern of power absorption followed by power generation paralleled the power profiles of the hip, ankle and MP joints. The total work performed by one hind limb was 0.71 J kg(-1). Surprisingly, the knee produced 85% of the work (0.60 J kg(-1)) done by the hind limb, and the positive work performed by the knee occurred during the first 40% of the take-off. There is little net work generated by the other three joints over the entire take-off. Velocity of the tuber coxae (a landmark on the pelvis of the animal) was negative (downward) during the first 40% of stance, which perhaps reflects the negative work performed to decrease the potential energy during the first 40% of contact. During the final 60% of contact, the hip, ankle and MP joints generate positive work, which is reflected in the increase of the animal's potential energy.

The effects of a single bout of downhill running and ensuing delayed onset of muscle soreness on running economy performed 48 h later.

Delayed onset of muscle soreness (DOMS) is a common response to exercise involving significant eccentric loading. Symptoms of DOMS vary widely and may include reduced force generating capacity, significant alterations in biochemical indices of muscle and connective tissue health, alteration of neuromuscular function, and changes in mechanical performance. The purpose of the investigation was to examine the effects of downhill running and ensuing DOMS on running economy and stride mechanics. Nine, well-trained distance runners and triathletes participated in the study. Running economy was measured at three relative intensities [65, 75, and 85% of maximal aerobic capacity ( VO(2peak))] before (RE1) and 48 h after (RE2) a 30-min downhill run (-10%) at 70% VO(2peak). Dependent variables included leg muscle soreness, rate of oxygen consumption ( VO(2)), minute ventilation, respiratory exchange ratio, lactate, heart rate, and stride length. These measurements were entered into a two-factor multivariate analysis of variance (MANOVA). The analysis revealed a significant time effect for all variables and a significant interaction (time x intensity) for lactate. The energy cost of locomotion was elevated at RE2 by an average of 3.2%. This was coupled with a significant reduction in stride length. The change in VO(2) was inversely correlated with the change in stride length ( r= -0.535). Lactate was significantly elevated at RE2 for each run intensity, with a mean increase of 0.61 mmol l(-1). Based on these findings, it is suggested that muscle damage led to changes in stride mechanics and a greater reliance on anaerobic methods of energy production, contributing to the change in running economy during DOMS.

Changes in spring-mass characteristics during treadmill running to exhaustion.

PURPOSE: To determine whether the stiffness characteristics of the leg change during a treadmill run to voluntary exhaustion. METHODS: Fifteen runners performed a test run at a constant speed that elicited approximately 80% of their .VO(2peak). The run was performed on a treadmill instrumented to measure vertical ground reaction forces; vertical stiffness and leg stiffness were calculated from these forces. Force data were sampled for 15 s every 5 min and immediately before the end of the test. From the force data, average stiffness characteristics were determined for each sample period. An ANOVA with repeated measures (alpha = 0.01) was performed for the group on both vertical and leg stiffness. A single-subject, case-series analysis was also performed on each subject by using ANOVA (alpha = 0.01). RESULTS: Group analysis revealed significant decreases (P < 0.01) in both vertical (23.9 to 23.1 kN.m(-1)) and leg (9.3 to 9.0 kN.m(-1)) stiffness over the run. Based on single-subject ANOVA, 14 of the 15 runners experienced significant (P < or = 0.01) changes in k(vert) over the run. A significant correlation between changes in stride rate and vertical stiffness was found (r = 0.85). Changes in the stiffness properties of the leg, as determined via the spring-mass model, resulted in changes in vertical displacement of the center of mass and leg length (distance from ankle to hip) during stance, as opposed to changes in peak force during ground contact. CONCLUSIONS: Observed changes in stride rate possibly result from changes in the stiffness characteristics of the leg during a run to fatigue.