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Biomedical subjects

R Kram

Publications and source records attributed to R Kram.

At least 19 recordsLinked to original sources
PubMed2001

Mechanical and metabolic determinants of the preferred step width in human walking.

We studied the selection of preferred step width in human walking by measuring mechanical and metabolic costs as a function of experimentally manipulated step width (0.00-0.45L, as a fraction of leg length L). We estimated mechanical costs from individual limb external mechanical work and metabolic costs using open circuit respirometry. The mechanical and metabolic costs both increased substantially (54 and 45%, respectively) for widths greater than the preferred value (0.15-0.45L) and with step width squared (R(2) = 0.91 and 0.83, respectively). As predicted by a three-dimensional model of walking mechanics, the increases in these costs appear to be a result of the mechanical work required for redirecting the centre of mass velocity during the transition between single stance phases (step-to-step transition costs). The metabolic cost for steps narrower than preferred (0.10-0.00L) increased by 8%, which was probably as a result of the added cost of moving the swing leg laterally in order to avoid the stance leg (lateral limb swing cost). Trade-offs between the step-to-step transition and lateral limb swing costs resulted in a minimum metabolic cost at a step width of 0.12L, which is not significantly different from foot width (0.11L) or the preferred step width (0.13L). Humans appear to prefer a step width that minimizes metabolic cost.

Biomechanical Phenomena↗
PubMed2001

Applied horizontal force increases impact loading in reduced-gravity running.

The chronic exposure of astronauts to microgravity results in structural degradation of their lower limb bones. Currently, no effective exercise countermeasure exists. On Earth, the impact loading that occurs with regular locomotion is associated with the maintenance of bone's structural integrity, but impact loads are rarely experienced in space. Accurately mimicking Earth-like impact loads in a reduced-gravity environment should help to reduce the degradation of bone caused by weightlessness. We previously showed that running with externally applied horizontal forces (AHF) in the anterior direction qualitatively simulates the high-impact loading associated with downhill running on Earth. We hypothesized that running with AHF at simulated reduced gravity would produce impact loads equal to or greater than values experienced during normal running at Earth gravity. With an AHF of 20% of gravity-specific body weight at all gravity levels, impact force peaks increased 74%, average impact loading rates increased 46%, and maximum impact loading rates increased 89% compared to running without any AHF. In contrast, AHF did not substantially affect active force peaks. Duty factor and stride frequency decreased modestly with AHF at all gravity levels. We found that running with an AHF in simulated reduced gravity produced impact loads equal to or greater than those experienced at Earth gravity. An appropriate AHF could easily augment existing partial gravity treadmill running exercise countermeasures used during spaceflight and help prevent musculoskeletal degradation.

Adult↗
PubMed2000

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↗
PubMed2000

Exploring dynamic similarity in human running using simulated reduced gravity.

The Froude number (a ratio of inertial to gravitational forces) predicts the occurrence of dynamic similarity in legged animals over a wide range of sizes and velocities for both walking and running gaits at Earth gravity. This is puzzling because the Froude number ignores elastic forces that are crucial for understanding running gaits. We used simulated reduced gravity as a tool for exploring dynamic similarity in human running. We simulated reduced gravity by applying a nearly constant upward force to the torsos of our subjects while they ran on a treadmill. We found that at equal Froude numbers, achieved through different combinations of velocity and levels of gravity, our subjects did not run in a dynamically similar manner. Thus, the inertial and gravitational forces that comprise the Froude number were not sufficient to characterize running in reduced gravity. Further, two dimensionless numbers that incorporate elastic forces, the Groucho number and the vertical Strouhal number, also failed to predict dynamic similarity in reduced-gravity running. To better understand the separate effects of velocity and gravity, we also studied running mechanics at fixed absolute velocities under different levels of gravity. The effects of velocity and gravity on the requirements of dynamic similarity differed in both magnitude and direction, indicating that there are no two velocity and gravity combinations at which humans will prefer to run in a dynamically similar manner. A comparison of walking and running results demonstrated that reduced gravity had different effects on the mechanics of each gait. This suggests that a single unifying hypothesis for the effects of size, velocity and gravity on both walking and running gaits will not be successful.

Animals↗
PubMed2000

The independent effects of gravity and inertia on running mechanics.

It is difficult to distinguish the independent effects of gravity from those of inertia on a running animal. Simply adding mass proportionally changes both the weight (gravitational force) and mass (inertial force) of the animal. We measured ground reaction forces for eight male humans running normally at 3 m s(-)(1) and under three experimental treatments: added gravitational and inertial forces, added inertial forces and reduced gravitational forces. Subjects ran at 110, 120 and 130 % of normal weight and mass, at 110, 120 and 130 % of normal mass while maintaining 100 % normal weight, and at 25, 50 and 75 % of normal weight while maintaining 100 % normal mass. The peak active vertical forces generated changed with weight, but did not change with mass. Surprisingly, horizontal impulses changed substantially more with weight than with mass. Gravity exerted a greater influence than inertia on both vertical and horizontal forces generated against the ground during running. Subjects changed vertical and horizontal forces proportionately at corresponding times in the step cycle to maintain the orientation of the resultant vector despite a nearly threefold change in magnitude across treatments. Maintaining the orientation of the resultant vector during periods of high force generation aligns the vector with the leg to minimize muscle forces.

Adult↗
PubMed2000

Muscular force or work: what determines the metabolic energy cost of running?

It has been proposed that the metabolic energy cost of running is determined by the magnitude and rate of muscular force generation needed to oppose gravity and operate muscle-tendon springs. Whole animal experiments, in vivo muscle force and fascicle length recordings, and in vitro muscle shortening velocity data support the idea.

Biomechanical Phenomena↗
PubMed1999

Walking in simulated reduced gravity: mechanical energy fluctuations and exchange.

Walking humans conserve mechanical and, presumably, metabolic energy with an inverted pendulum-like exchange of gravitational potential energy and horizontal kinetic energy. Walking in simulated reduced gravity involves a relatively high metabolic cost, suggesting that the inverted-pendulum mechanism is disrupted because of a mismatch of potential and kinetic energy. We tested this hypothesis by measuring the fluctuations and exchange of mechanical energy of the center of mass at different combinations of velocity and simulated reduced gravity. Subjects walked with smaller fluctuations in horizontal velocity in lower gravity, such that the ratio of horizontal kinetic to gravitational potential energy fluctuations remained constant over a fourfold change in gravity. The amount of exchange, or percent recovery, at 1.00 m/s was not significantly different at 1.00, 0.75, and 0.50 G (average 64.4%), although it decreased to 48% at 0.25 G. As a result, the amount of work performed on the center of mass does not explain the relatively high metabolic cost of walking in simulated reduced gravity.

Aged↗
PubMed1999

Metabolic cost of generating horizontal forces during human running.

Previous studies have suggested that generating vertical force on the ground to support body weight (BWt) is the major determinant of the metabolic cost of running. Because horizontal forces exerted on the ground are often an order of magnitude smaller than vertical forces, some have reasoned that they have negligible cost. Using applied horizontal forces (AHF; negative is impeding, positive is aiding) equal to -6, -3, 0, +3, +6, +9, +12, and +15% of BWt, we estimated the cost of generating horizontal forces while subjects were running at 3.3 m/s. We measured rates of oxygen consumption (VO2) for eight subjects. We then used a force-measuring treadmill to measure ground reaction forces from another eight subjects. With an AHF of -6% BWt, VO2 increased 30% compared with normal running, presumably because of the extra work involved. With an AHF of +15% BWt, the subjects exerted approximately 70% less propulsive impulse and exhibited a 33% reduction in VO2. Our data suggest that generating horizontal propulsive forces constitutes more than one-third of the total metabolic cost of normal running.

Adult↗
PubMed1998

Energetics and biomechanics of locomotion by red kangaroos (Macropus rufus).

As red kangaroos hop faster over level ground, their rate of oxygen consumption (indicating metabolic energy consumption) remains nearly the same. This phenomenon has been attributed to exceptional elastic energy storage and recovery via long compliant tendons in the legs. Alternatively, red kangaroos may have exceptionally efficient muscles. To estimate efficiency, we measured the metabolic cost of uphill hopping, where muscle fibers must perform mechanical work against gravity. We found that uphill hopping was much more expensive than level hopping. The maximal rate of oxygen consumption measured (3 ml O2 kg-1 s-1) exceeds all but a few vertebrate species. However, efficiency values were normal, approximately 30%. At faster level hopping speeds the effective mechanical advantage of the extensor muscles of the ankle joint remained the same. Thus, kangaroos generate the same muscular force at all speeds but do so more rapidly at faster hopping speeds. This contradicts a recent hypothesis for what sets the cost of locomotion. The cost of transport (J kg-1 m-1) decreases at faster hopping speeds, yet red kangaroos prefer to use relatively slow speeds that avoid high levels of tendon stress.

Animals↗
PubMed1998

Force treadmill for measuring vertical and horizontal ground reaction forces.

We constructed a force treadmill to measure the vertical, horizontal and lateral components of the ground-reaction forces (Fz, Fy, Fx, respectively) and the ground-reaction force moments (Mz, My, Mx), respectively exerted by walking and running humans. The chassis of a custom-built, lightweight (90 kg), mechanically stiff treadmill was supported along its length by a large commercial force platform. The natural frequencies of vibration were >178 Hz for Fz and >87 Hz for Fy, i.e., well above the signal content of these ground-reaction forces. Mechanical tests and comparisons with data obtained from a force platform runway indicated that the force treadmill recorded Fz, Fy, Mx and My ground-reaction forces and moments accurately. Although the lowest natural frequency of vibration was 88 Hz for Fx, the signal-to-noise ratios for Fx and Mz were unacceptable. This device greatly decreases the time and laboratory space required for locomotion experiments and clinical evaluations. The modular design allows for independent use of both treadmill and force platform.

Biophysical Phenomena↗
PubMed1998

Energetics of bipedal running. I. Metabolic cost of generating force.

Similarly sized bipeds and quadrupeds use nearly the same amount of metabolic energy to run, despite dramatic differences in morphology and running mechanics. It has been shown that the rate of metabolic energy use in quadrupedal runners and bipedal hoppers can be predicted from just body weight and the time available to generate force as indicated by the duration of foot-ground contact. We tested whether this link between running mechanics and energetics also applies to running bipeds. We measured rates of energy consumption and times of foot contact for humans (mean body mass 78.88 kg) and five species of birds (mean body mass range 0.13-40.1 kg). We find that most (70-90%) of the increase in metabolic rate with speed in running bipeds can be explained by changes in the time available to generate force. The rate of force generation also explains differences in metabolic rate over the size range of birds measured. However, for a given rate of force generation, birds use on average 1.7 times more metabolic energy than quadrupeds. The rate of energy consumption for a given rate of force generation for humans is intermediate between that of birds and quadrupeds. These results support the idea that the cost of muscular force production determines the energy cost of running and suggest that bipedal runners use more energy for a given rate of force production because they require a greater volume of muscle to support their body weight.

Animals↗
PubMed1997

Three-dimensional kinematics and limb kinetic energy of running cockroaches.

We tested the hypothesis that fast-running hexapeds must generate high levels of kinetic energy to cycle their limbs rapidly compared with bipeds and quadrupeds. We used high-speed video analysis to determine the three-dimensional movements of the limbs and bodies of cockroaches (Blaberus discoidalis) running on a motorized treadmill at 21 cm s-1 using an alternating tripod gait. We combined these kinematic data with morphological data to calculate the mechanical energy produced to move the limbs relative to the overall center of mass and the mechanical energy generated to rotate the body (head + thorax + abdomen) about the overall center of mass. The kinetic energy involved in moving the limbs was 8 microJ stride-1 (a power output of 21 mW kg-1, which was only approximately 13% of the external mechanical energy generated to lift and accelerate the overall center of mass at this speed. Pitch, yaw and roll rotational movements of the body were modest (less than +/- 7 degrees), and the mechanical energy required for these rotations was surprisingly small (1.7 microJ stride-1 for pitch, 0.5 microJ stride-1 for yaw and 0.4 microJ stride-1 for roll) as was the power (4.2, 1.2 and 1.1 mW kg-1, respectively). Compared at the same absolute forward speed, the mass-specific kinetic energy generated by the trotting hexaped to swing its limbs was approximately half of that predicted from data on much larger two- and four-legged animals. Compared at an equivalent speed (mid-trotting speed), limb kinetic energy was a smaller fraction of total mechanical energy for cockroaches than for large bipedal runners and hoppers and for quadrupedal trotters. Cockroaches operate at relatively high stride frequencies, but distribute ground reaction forces over a greater number of relatively small legs. The relatively small leg mass and inertia of hexapeds may allow relatively high leg cycling frequencies without exceptionally high internal mechanical energy generation.

Animals↗
PubMed1997

The effect of reduced gravity on the kinematics of human walking: a test of the dynamic similarity hypothesis for locomotion.

To gain insight into the basic principles that govern the biomechanics of locomotion, we investigated the effect of reduced gravity on walking kinematics. We hypothesized that humans walk in a dynamically similar fashion at combinations of speed and simulated gravity that provide equal values of the Froude number, v2/gLleg, where v is forward speed, g is gravitational acceleration and Lleg is leg length. The Froude number has been used to predict the kinematics and kinetics of legged locomotion over a wide range of animal sizes and speeds, and thus provides a potentially unifying theory for the combined effects of speed, size and gravity on locomotion biomechanics. The occurrence of dynamic similarity at equal Froude numbers has been attributed previously to the importance of gravitational forces in determining locomotion mechanics. We simulated reduced gravity using a device that applies a nearly constant upward force to the torso while subjects walked on a treadmill. We found that at equal Froude numbers, under different levels of gravity (0.25g-1.0g), the subjects walked with nearly the same duty factor (ratio of contact time to stride time), but with relative stride lengths (Ls/Lleg, where Ls is stride length) that differed by as much as 67 %, resulting in the rejection of our hypothesis. To understand the separate effects of speed and gravity further, we compared the mechanics of walking at the same absolute speed at different levels of gravity (0.25g-1.0g). In lower gravity, subjects walked with lower duty factors (10 %) and shorter relative stride lengths (16 %). These modest changes in response to the fourfold change in gravity indicate that factors other than gravitational forces are the primary determinants of walking biomechanics.

Biomechanical Phenomena↗
PubMed1997

Effect of reduced gravity on the preferred walk-run transition speed.

We investigated the effect of reduced gravity on the human walk-run gait transition speed and interpreted the results using an inverted-pendulum mechanical model. We simulated reduced gravity using an apparatus that applied a nearly constant upward force at the center of mass, and the subjects walked and ran on a motorized treadmill. In the inverted pendulum model for walking, gravity provides the centripetal force needed to keep the pendulum in contact with the ground. The ratio of the centripetal and gravitational forces (mv2/L)/(mg) reduces to the dimensionless Froude number (v2/gL). Applying this model to a walking human, m is body mass, v is forward velocity, L is leg length and g is gravity. In normal gravity, humans and other bipeds with different leg lengths all choose to switch from a walk to a run at different absolute speeds but at approximately the same Froude number (0.5). We found that, at lower levels of gravity, the walk-run transition occurred at progressively slower absolute speeds but at approximately the same Froude number. This supports the hypothesis that the walk-run transition is triggered by the dynamics of an inverted-pendulum system.

Energy Metabolism↗
PubMed1996

Musculoskeletal adaptations to weightlessness and development of effective countermeasures.

A Research Roundtable, organized by the American College of Sports Medicine with sponsorship from the National Aeronautics and Space Administration, met in November 1995 to define research strategies for effective exercise countermeasures to weightlessness. Exercise was considered both independently of, and in conjunction with, other therapeutic modalities (e.g., pharmacological nutritional, hormonal, and growth-related factors) that could prevent or minimize the structural and functional deficits involving skeletal muscle and bone in response to chronic exposure to weightlessness, as well as return to Earth baseline function if a degree of loss is inevitable. Musculoskeletal deficits and countermeasures are described with respect to: 1) muscle and connective tissue atrophy and localized bone loss, 2) reductions in motor performance, 3) potential proneness to injury of hard and soft tissues, and 4) probable interaction between muscle atrophy and cardiovascular alterations that contribute to the postural hypotension observed immediately upon return from space flight. In spite of a variety of countermeasure protocols utilized previously involving largely endurance types of exercise, there is presently no activity-specific countermeasure(s) that adequately prevent or reduce musculoskeletal deficiencies. It seems apparent that countermeasure exercises that have a greater resistance element, as compared to endurance activities, may prove beneficial to the musculoskeletal system. Many questions remain for scientific investigation to identify efficacious countermeasure protocols, which will be imperative with the emerging era of long-term space flight.

Adaptation, Physiological↗
PubMed1996

Inexpensive load carrying by rhinoceros beetles

These experiments determined the magnitude of loads that rhinoceros beetles (Scarabaeidae) can carry and also the metabolic energy required for carrying loads. I hypothesized that, like many other animals, these beetles would have metabolic rates in direct proportion to the total load (body mass plus added mass). Eight beetles (Xylorctes thestalus) walked at 1 cm s-1 on a motorized treadmill enclosed in a respirometer. The beetles could sustain this speed with loads of more than 30 times their body mass. In addition to being strong, these beetles carry loads with remarkable economy. The metabolic cost of moving a gram of additional load was more than five times cheaper than that of moving a gram of body mass. This phenomenon cannot be explained by conventional models that link the biomechanics and metabolic energy cost of locomotion.

Journal Article↗
PubMed1995

Moving cheaply: energetics of walking in the African elephant.

Large animals have a much better fuel economy than small ones, both when they rest and when they run. At rest, each gram of tissue of the largest land animal, the African elephant, consumes metabolic energy at 1/20 the rate of a mouse; using existing allometric relationships, we calculate that it should be able to carry 1 g of its tissue (or a load) for 1 km at 1/40 the cost for a mouse. These relationships between energetics and size are so consistent that they have been characterized as biological laws. The elephant has massive legs and lumbers along awkwardly, suggesting that it might expend more energy to move about than other animals. We find, however, that its energetic cost of locomotion is predicted remarkably well by the allometric relationships and is the lowest recorded for any living land animal.

Animals↗