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Biomedical subjects

C T Farley

Publications and source records attributed to C T Farley.

At least 19 recordsLinked to original sources

Soleus H-reflex gain in humans walking and running under simulated reduced gravity.

The Hoffmann (H-) reflex is an electrical analogue of the monosynaptic stretch reflex, elicited by bypassing the muscle spindle and directly stimulating the afferent nerve. Studying H-reflex modulation provides insight into how the nervous system centrally modulates stretch reflex responses.A common measure of H-reflex gain is the slope of the relationship between H-reflex amplitude and EMG amplitude. To examine soleus H-reflex gain across a range of EMG levels during human locomotion, we used simulated reduced gravity to reduce muscle activity. We hypothesised that H-reflex gain would be independent of gravity level.We recorded EMG from eight subjects walking (1.25 m s-1) and running (3.0 m s-1) at four gravity levels (1.0, 0.75, 0.5 and 0.25 G (Earth gravity)). We normalised the stimulus M-wave and resulting H-reflex to the maximal M-wave amplitude (Mmax) elicited throughout the stride to correct for movement of stimulus and recording electrodes relative to nerve and muscle fibres. Peak soleus EMG amplitude decreased by ~30% for walking and for running over the fourfold change in gravity. As hypothesised, slopes of linear regressions fitted to H-reflex versus EMG data were independent of gravity for walking and running (ANOVA, P > 0.8). The slopes were also independent of gait (P > 0.6), contrary to previous studies. Walking had a greater y-intercept (19.9% Mmax) than running (-2.5% Mmax; P < 0.001). At all levels of EMG, walking H-reflex amplitudes were higher than running H-reflex amplitudes by a constant amount. We conclude that the nervous system adjusts H-reflex threshold but not H-reflex gain between walking and running. These findings provide insight into potential neural mechanisms responsible for spinal modulation of the stretch reflex during human locomotion.

Adult↗

How animals move: an integrative view.

Recent advances in integrative studies of locomotion have revealed several general principles. Energy storage and exchange mechanisms discovered in walking and running bipeds apply to multilegged locomotion and even to flying and swimming. Nonpropulsive lateral forces can be sizable, but they may benefit stability, maneuverability, or other criteria that become apparent in natural environments. Locomotor control systems combine rapid mechanical preflexes with multimodal sensory feedback and feedforward commands. Muscles have a surprising variety of functions in locomotion, serving as motors, brakes, springs, and struts. Integrative approaches reveal not only how each component within a locomotor system operates but how they function as a collective whole.

Animals↗

Leg stiffness primarily depends on ankle stiffness during human hopping.

When humans hop in place or run forward, they adjust leg stiffness to accommodate changes in stride frequency or surface stiffness. The goal of the present study was to determine the mechanisms by which humans adjust leg stiffness during hopping in place. Five subjects hopped in place at 2.2 Hz while we collected force platform and kinematic data. Each subject completed trials in which they hopped to whatever height they chose ("preferred height hopping") and trials in which they hopped as high as possible ("maximum height hopping"). Leg stiffness was approximately twice as great for maximum height hopping as for preferred height hopping. Ankle torsional stiffness was 1.9-times greater while knee torsional stiffness was 1.7-times greater in maximum height hopping than in preferred height hopping. We used a computer simulation to examine the sensitivity of leg stiffness to the observed changes in ankle and knee stiffness. Our model consisted of four segments (foot, shank, thigh, head-arms-trunk) interconnected by three torsional springs (ankle, knee, hip). In the model, increasing ankle stiffness by 1.9-fold, as observed in the subjects, caused leg stiffness to increase by 2.0-fold. Increasing knee stiffness by 1.7-fold had virtually no effect on leg stiffness. Thus, we conclude that the primary mechanism for leg stiffness adjustment is the adjustment of ankle stiffness.

Adult↗

Runners adjust leg stiffness for their first step on a new running surface.

Human runners adjust the stiffness of their stance leg to accommodate surface stiffness during steady state running. This adjustment allows runners to maintain similar center of mass movement (e.g., ground contact time and stride frequency) regardless of surface stiffness. When runners encounter abrupt transitions in the running surface, they must either make a rapid adjustment or allow the change in the surface stiffness to disrupt their running mechanics. Our goal was to determine how quickly runners adjust leg stiffness when they encounter an abrupt but expected change in surface stiffness that they have encountered previously. Six human subjects ran at 3 m s(-1) on a rubber track with two types of rubber surfaces: a compliant "soft" surface (ksurf = 21.3 kN m(-1) and a non-compliant "hard" surface (ksurf = 533 kN m(-1). We found that runners completely adjusted leg stiffness for their first step on the new surface after the transition. For example, runners decreased leg stiffness by 29% between the last step on the soft surface and the first step on the hard surface (from 10.7 kN m(-1) to 7.6 kN m(-1), respectively). As a result, the vertical displacement of the center of mass during stance ( approximately 7 cm) did not change at the transition despite a reduction in surface compression from 6 cm to less than 0.25 cm. By rapidly adjusting leg stiffness, each runner made a smooth transition between surfaces so that the path of the center of mass was unaffected by the change in surface stiffness.

Adult↗

Running in the real world: adjusting leg stiffness for different surfaces.

A running animal coordinates the actions of many muscles, tendons, and ligaments in its leg so that the overall leg behaves like a single mechanical spring during ground contact. Experimental observations have revealed that an animal's leg stiffness is independent of both speed and gravity level, suggesting that it is dictated by inherent musculoskeletal properties. However, if leg stiffness was invariant, the biomechanics of running (e.g. peak ground reaction force and ground contact time) would change when an animal encountered different surfaces in the natural world. We found that human runners adjust their leg stiffness to accommodate changes in surface stiffness, allowing them to maintain similar running mechanics on different surfaces. These results provide important insight into mechanics and control of animal locomotion and suggest that incorporating an adjustable leg stiffness in the design of hopping and running robots is important if they are to match the agility and speed of animals on varied terrain.

Animals↗

Mechanism of leg stiffness adjustment for hopping on surfaces of different stiffnesses.

When humans hop in place or run forward, leg stiffness is increased to offset reductions in surface stiffness, allowing the global kinematics and mechanics to remain the same on all surfaces. The purpose of the present study was to determine the mechanism for adjusting leg stiffness. Seven subjects hopped in place on surfaces of different stiffnesses (23-35,000 kN/m) while force platform, kinematic, and electromyographic data were collected. Leg stiffness approximately doubled between the most stiff surface and the least stiff surface. Over the same range of surfaces, ankle torsional stiffness increased 1.75-fold, and the knee became more extended at the time of touchdown (2.81 vs. 2.65 rad). We used a computer simulation to examine the sensitivity of leg stiffness to the observed changes in ankle stiffness and touchdown knee angle. Our model consisted of four segments (foot, shank, thigh, head-arms-trunk) interconnected by three torsional springs (ankle, knee, hip). In the model, an increase in ankle stiffness 1.75-fold caused leg stiffness to increase 1.7-fold. A change in touchdown knee angle as observed in the subjects caused leg stiffness to increase 1.3-fold. Thus both joint stiffness and limb geometry adjustments are important in adjusting leg stiffness to allow similar hopping on different surfaces.

Adult↗

Determinants of the center of mass trajectory in human walking and running.

Walking is often modeled as an inverted pendulum system in which the center of mass vaults over the rigid stance limb. Running is modeled as a simple spring-mass system in which the center of mass bounces along on the compliant stance limb. In these models, differences in stance-limb behavior lead to nearly opposite patterns of vertical movements of the center of mass in the two gaits. Our goal was to quantify the importance of stance-limb behavior and other factors in determining the trajectory of the center of mass during walking and running. We collected kinematic and force platform data during human walking and running. Virtual stance-limb compression (i.e. reduction in the distance between the point of foot-ground contact and the center of mass during the first half of the stance phase) was only 26% lower for walking (0.091 m) than for running (0.123 m) at speeds near the gait transition speed. In spite of this relatively small difference, the center of mass moved upwards by 0.031 m during the first half of the stance phase during walking and moved downwards by 0.073 m during the first half of the stance phase during running. The most important reason for this difference was that the stance limb swept through a larger angle during walking (30.4 degrees) than during running (19.2 degrees). We conclude that stance-limb touchdown angle and virtual stance-limb compression both play important roles in determining the trajectory of the center of mass and whether a gait is a walk or a run.

Biomechanical Phenomena↗

Interaction of leg stiffness and surfaces stiffness during human hopping.

When mammals run, the overall musculoskeletal system behaves as a single linear "leg spring". We used force platform and kinematic measurements to determine whether leg spring stiffness (k(leg)) is adjusted to accommodate changes in surface stiffness (ksurf) when humans hoop in place, a good experimental model for examining adjustments to k(leg) in bouncing gaits. We found that k(leg) was greatly increased to accommodate surfaces of lower stiffnesses. The series combination of k(leg) and ksurf [total stiffness (ktot)] was independent of ksurf at a given hopping frequency. For example, when humans hopped at a frequency of 2 Hz, they tripled their k(leg) on the least stiff surface (ksurf = 26.1 kN/m; k(leg) = 53.3 kN/m) compared with the most stiff surface (ksurf = 35,000 kN/m; k(leg) = 17.8 kN/m). Values for ktot were not significantly different on the least stiff surface (16.7 kN/m) and the most stiff surface (17.8 kN/m). Because of the k(leg) adjustment, many aspects of the hopping mechanics (e.g., ground-contact time and center of mass vertical displacement) remained remarkably similar despite a > 1,000-fold change in ksurf. This study provides insight into how k(leg) adjustments can allow similar locomotion mechanics on the variety of terrains encountered by runners in the natural world.

Aged↗

Mechanics of locomotion in lizards.

Lizards bend their trunks laterally with each step of locomotion and, as a result, their locomotion appears to be fundamentally different from mammalian locomotion. The goal of the present study was to determine whether lizards use the same two basic gaits as other legged animals or whether they use a mechanically unique gait due to lateral trunk bending. Force platform and kinematic measurements revealed that two species of lizards, Coleonyx variegatus and Eumeces skiltonianus, used two basic gaits similar to mammalian walking and trotting gaits. In both gaits, the kinetic energy fluctuations due to lateral movements of the center of mass were less than 5% of the total external mechanical energy fluctuations. In the walking gait, both species vaulted over their stance limbs like inverted pendulums. The fluctuations in kinetic energy and gravitational potential energy of the center of mass were approximately 180 degrees out of phase. The lizards conserved as much as 51% of the external mechanical energy required for locomotion by the inverted pendulum mechanism. Both species also used a bouncing gait, similar to mammalian trotting, in which the fluctuations in kinetic energy and gravitational potential energy of the center of mass were nearly exactly in phase. The mass-specific external mechanical work required to travel 1 m (1.5 J kg-1) was similar to that for other legged animals. Thus, in spite of marked lateral bending of the trunk, the mechanics of lizard locomotion is similar to the mechanics of locomotion in other legged animals.

Animals↗

Maximum speed and mechanical power output in lizards.

The goal of the present study was to test the hypothesis that maximum running speed is limited by how much mechanical power the muscular system can produce. To test this hypothesis, two species of lizards, Coleonyx variegatus and Eumeces skiltonianus, sprinted on hills of different slopes. According to the hypothesis, maximum speed should decrease on steeper uphill slopes but mechanical power output at maximum speed should be independent of slope. For level sprinting, the external mechanical power output was determined from force platform data. For uphill sprinting, the mechanical power output was approximated as the power required to lift the center of mass vertically. When the slope increased from level to 40 degrees uphill, maximum speed decreased by 28% in C. variegatus and by 16% in E. skiltonianus. At maximum speed on a 40 degrees uphill slope in both species, the mechanical power required to lift the body vertically was approximately 3.9 times greater than the external mechanical power output at maximum speed on the level. Because total limb mass is small in both species (6-16% of body mass) and stride frequency is similar at maximum speed on all slopes, the internal mechanical power output is likely to be small and similar in magnitude on all slopes. I conclude that the muscular system is capable of producing substantially more power during locomotion than it actually produces during level sprinting. Thus, the capacity of the muscular system to produce power does not limit maximum running speed.

Animals↗

Leg stiffness and stride frequency in human running.

When humans and other mammals run, the body's complex system of muscle, tendon and ligament springs behaves like a single linear spring ('leg spring'). A simple spring-mass model, consisting of a single linear leg spring and a mass equivalent to the animal's mass, has been shown to describe the mechanics of running remarkably well. Force platform measurements from running animals, including humans, have shown that the stiffness of the leg spring remains nearly the same at all speeds and that the spring-mass system is adjusted for higher speeds by increasing the angle swept by the leg spring. The goal of the present study is to determine the relative importance of changes to the leg spring stiffness and the angle swept by the leg spring when humans alter their stride frequency at a given running speed. Human subjects ran on treadmill-mounted force platform at 2.5ms-1 while using a range of stride frequencies from 26% below to 36% above the preferred stride frequency. Force platform measurements revealed that the stiffness of the leg spring increased by 2.3-fold from 7.0 to 16.3 kNm-1 between the lowest and highest stride frequencies. The angle swept by the leg spring decreased at higher stride frequencies, partially offsetting the effect of the increased leg spring stiffness on the mechanical behavior of the spring-mass system. We conclude that the most important adjustment to the body's spring system to accommodate higher stride frequencies is that leg spring becomes stiffer.

Adaptation, Physiological↗

Running springs: speed and animal size.

Trotting and hopping animals use muscles, tendons and ligaments to store and return elastic energy as they bounce along the ground. We examine how the musculoskeletal spring system operates at different speeds and in animals of different sizes. We model trotting and hopping as a simple spring-mass system which consists of a leg spring and a mass. We find that the stiffness of the leg spring (k(leg)) is nearly independent of speed in dogs, goats, horses and red kangaroos. As these animals trot or hop faster, the leg spring sweeps a greater angle during the stance phase, and the vertical excursion of the center of mass during the ground contact phase decreases. The combination of these changes to the spring system causes animals to bounce off the ground more quickly at higher speeds. Analysis of a wide size range of animals (0.1-140 kg) at equivalent speeds reveals that larger animals have stiffer leg springs (k(leg) [symbol: see text] M0.67, where M is body mass), but that the angle swept by the leg spring is nearly independent of body mass. As a result, the resonant period of vertical vibration of the spring-mass system is longer in larger animals. The length of time that the feet are in contact with the ground increases with body mass in nearly the same way as the resonant period of vertical vibration.

Animals↗

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.

Energy Metabolism↗

A mechanical trigger for the trot-gallop transition in horses.

It is widely thought that animals switch gaits at speeds that minimize energetic cost. Horses naturally switched from a trot to a gallop at a speed where galloping required more energy than trotting, and thus, the gait transition actually increased the energetic cost of running. However, by galloping at this speed, the peak forces on the muscles, tendons, and bones, and presumably the chance of injury, are reduced. When the horses carried weights, they switched from a trot to a gallop at a lower speed but at the same critical level of force. These findings suggest that the trot-gallop transition is triggered when musculoskeletal forces reach a critical level.

Animals↗

Hopping frequency in humans: a test of how springs set stride frequency in bouncing gaits.

The storage and recovery of elastic energy in muscle-tendon springs is important in running, hopping, trotting, and galloping. We hypothesized that animals select the stride frequency at which they behave most like simple spring-mass systems. If higher or lower frequencies are used, they will not behave like simple spring-mass systems, and the storage and recovery of elastic energy will be reduced. We tested the hypothesis by having humans hop forward on a treadmill over a range of speeds and hop in place over a range of frequencies. The body was modeled as a simple spring-mass system, and the properties of the spring were measured by use of a force platform. Our subjects used nearly the same frequency (the "preferred frequency," 2.2 hops/s) when they hopped forward on a treadmill and when they hopped in place. At this frequency, the body behaved like a simple spring-mass system. Contrary to our predictions, it also behaved like a simple spring-mass system when the subjects hopped at higher frequencies, up to the maximum they could achieve. However, at the higher frequencies, the time available to apply force to the ground (the ground contact time) was shorter, perhaps resulting in a higher cost of generating muscular force. At frequencies below the preferred frequency, as predicted by the hypothesis, the body did not behave in a springlike manner, and it appeared likely that the storage and recovery of elastic energy was reduced. The combination of springlike behavior and a long ground contact time at the preferred frequency should minimize the cost of generating muscular force.

Adult↗