PubMed Health⌕ Search

Biomedical subjects

Rodger Kram

Publications and source records attributed to Rodger Kram.

16 recordsLinked to original sources
PubMed2006

The locomotor kinematics of Asian and African elephants: changes with speed and size.

For centuries, elephant locomotion has been a contentious and confusing challenge for locomotion scientists to understand, not only because of technical difficulties but also because elephant locomotion is in some ways atypical of more familiar quadrupedal gaits. We analyzed the locomotor kinematics of over 2400 strides from 14 African and 48 Asian elephant individuals (body mass 116-4632 kg) freely moving over ground at a 17-fold range of speeds, from slow walking at 0.40 m s(-1) to the fastest reliably recorded speed for elephants, 6.8 m s(-1). These data reveal that African and Asian elephants have some subtle differences in how size-independent kinematic parameters change with speed. Although elephants use a lateral sequence footfall pattern, like many other quadrupeds, they maintain this footfall pattern at all speeds, shifting toward a 25% phase offset between limbs (singlefoot) as they increase speed. The duty factors of elephants are greater for the forelimbs than for the hindlimbs, so an aerial phase for the hindquarters is reached at slower speeds than for the forequarters. This aerial phase occurs at a Froude number of around 1, matching theoretical predictions. At faster speeds, stance and swing phase durations approach asymptotes, with the duty factor beginning to level off, concurrent with an increase in limb compliance that likely keeps peak forces relatively low. This increase of limb compliance is reflected by increased compression of the hindlimbs. Like other tetrapods, smaller elephants are relatively more athletic than larger ones, but still move very similarly to adults even at <500 kg. At any particular speed they adopt greater relative stride frequencies and relative stride lengths compared to larger elephants. This extends to near-maximal locomotor performance as well - smaller elephants reach greater Froude numbers and smaller duty factors, hence likely reach relatively greater peak loads on their limbs and produce this force more rapidly. A variety of lines of kinematic evidence support the inference that elephants change their mechanics near a Froude number of 1 (if not at slower speeds), at least to using more compliant limbs, if not spring-like whole-body kinetics. In some ways, elephants move similarly to many other quadrupeds, such as increasing speed mainly by increasing stride frequency (except at fast speeds), and they match scaling predictions for many stride parameters. The main difference from most other animals is that elephants never change their footfall pattern to a gait that uses a whole-body aerial phase. Our large dataset establishes what the normal kinematics of elephant locomotion are, and can also be applied to identify gait abnormalities that may signal musculoskeletal pathologies, a matter of great importance to keepers of captive elephants.

Animals↗
PubMed2006

Mechanical energy fluctuations during hill walking: the effects of slope on inverted pendulum exchange.

Humans and other animals exchange gravitational potential energy (GPE) and kinetic energy (KE) of the center of mass during level walking. How effective is this energy exchange during downhill and uphill walking? Based on previous reports and our own reasoning, we expected that during downhill walking, the possibility for mechanical energy exchange would be enhanced and during uphill walking, the possibility for exchange would be reduced. We measured the fluctuations of the mechanical energies for five men and five women walking at 1.25 m s(-1). Subjects walked on the level, downhill, and uphill on a force measuring treadmill mounted at 3 degrees, 6 degrees and 9 degrees. We evaluated energy exchange during the single support period based on the GPE and KE fluctuation factors of phase relationship, relative magnitude and extent of symmetry. As expected, during level walking, the GPE and KE curves were out of phase, of similar magnitude, and nearly mirror images so that the fluctuations in combined (GPE+KE) energy were attenuated. During downhill walking, the fluctuations in the combined energy of the center of mass were smaller than those on the level, i.e. mechanical energy exchange was more effective. During uphill walking, the fluctuations in the combined energy of the center of mass were larger than those on the level, i.e. mechanical energy exchange was less effective. Mechanical energy exchange occurred during downhill, level and uphill walking, but it was most effective during downhill walking.

Adult↗
PubMed2005

Effects of obesity and sex on the energetic cost and preferred speed of walking.

The metabolic energy cost of walking is determined, to a large degree, by body mass, but it is not clear how body composition and mass distribution influence this cost. We tested the hypothesis that walking would be most expensive for obese women compared with obese men and normal-weight women and men. Furthermore, we hypothesized that for all groups, preferred walking speed would correspond to the speed that minimized the gross energy cost per distance. We measured body composition, maximal oxygen consumption, and preferred walking speed of 39 (19 class II obese, 20 normal weight) women and men. We also measured oxygen consumption and carbon dioxide production while the subjects walked on a level treadmill at six speeds (0.50-1.75 m/s). Both obesity and sex affected the net metabolic rate (W/kg) of walking. Net metabolic rates of obese subjects were only approximately 10% greater (per kg) than for normal-weight subjects, and net metabolic rates for women were approximately 10% greater than for men. The increase in net metabolic rate at faster walking speeds was greatest in obese women compared with the other groups. Preferred walking speed was not different across groups (1.42 m/s) and was near the speed that minimized gross energy cost per distance. Surprisingly, mass distribution (thigh mass/body mass) was not related to net metabolic rate, but body composition (% fat) was (r2= 0.43). Detailed biomechanical studies of walking are needed to investigate whether obese individuals adopt novel energy saving mechanisms during walking.

Adult↗
PubMed2005

Ground reaction forces during downhill and uphill running.

We investigated the normal and parallel ground reaction forces during downhill and uphill running. Our rationale was that these force data would aid in the understanding of hill running injuries and energetics. Based on a simple spring-mass model, we hypothesized that the normal force peaks, both impact and active, would increase during downhill running and decrease during uphill running. We anticipated that the parallel braking force peaks would increase during downhill running and the parallel propulsive force peaks would increase during uphill running. But, we could not predict the magnitude of these changes. Five male and five female subjects ran at 3m/s on a force treadmill mounted on the level and on 3 degrees, 6 degrees, and 9 degrees wedges. During downhill running, normal impact force peaks and parallel braking force peaks were larger compared to the level. At -9 degrees, the normal impact force peaks increased by 54%, and the parallel braking force peaks increased by 73%. During uphill running, normal impact force peaks were smaller and parallel propulsive force peaks were larger compared to the level. At +9 degrees, normal impact force peaks were absent, and parallel propulsive peaks increased by 75%. Neither downhill nor uphill running affected normal active force peaks. Combined with previous biomechanics studies, our normal impact force data suggest that downhill running substantially increases the probability of overuse running injury. Our parallel force data provide insight into past energetic studies, which show that the metabolic cost increases during downhill running at steep angles.

Adult↗
PubMed2005

Energetic cost and preferred speed of walking in obese vs. normal weight women.

OBJECTIVE: We tested the hypotheses that walking is more expensive for obese women, and they prefer slower walking speeds that minimize the gross energy cost per distance despite a greater relative aerobic effort [percent of maximal oxygen uptake (Vo(2max))/kg]. RESEARCH METHODS AND PROCEDURES: Twenty adult women, 10 obese (BMI = 34.1 +/- 3.2 kg/m(2)) and 10 normal weight (BMI = 20.4 +/- 2.1 kg/m(2)) volunteered. To determine the metabolic rate and energy cost per distance vs. speed relationships, we measured Vo(2) and V(CO(2)) while subjects walked on a treadmill at six speeds (0.50, 0.75, 1.0, 1.25, 1.5, and 1.75 m/s; 5-minute trials, with a 5-minute rest period between trials). We measured preferred walking speed on a 50-m section of level sidewalk and Vo(2max) using a modified Balke treadmill protocol. RESULTS: Walking was 11% more expensive for the obese subjects, but they preferred to walk at similar speeds as normal weight subjects (1.40 vs. 1.47 m/s, p = 0.07). Both groups preferred walking speeds at which their gross energy cost per distance was almost minimized. Obese subjects had a smaller Vo(2max)/kg, so they required a greater relative aerobic effort at the preferred speed (51% vs. 36%, p = 0.001). DISCUSSION: Obese women preferred a walking speed that minimized energy cost per distance, even though this strategy required a greater relative aerobic effort than walking more slowly. Our results suggest that walking slower for a set distance may be an appropriate exercise recommendation for a weight management prescription in obese adults.

Adult↗
PubMed2005

Metabolic energy and muscular activity required for leg swing in running.

The metabolic cost of leg swing in running is highly controversial. We investigated the cost of initiating and propagating leg swing at a moderate running speed and some of the muscular actions involved. We constructed an external swing assist (ESA) device that applied small anterior pulling forces to each foot during the first part of the swing phase. Subjects ran on a treadmill at 3.0 m/s normally and with ESA forces up to 4% body weight. With the greatest ESA force, net metabolic rate was 20.5% less than during normal running. Thus we infer that the metabolic cost of initiating and propagating leg swing comprises approximately 20% of the net cost of normal running. Even with the greatest ESA, mean electromyograph (mEMG) of the medial gastrocnemius and soleus muscles during later portions of stance phase did not change significantly compared with normal running, indicating that these muscles are not responsible for the initiation of leg swing. However, with ESA, rectus femoris mEMG during the early portions of swing phase was as much as 74% less than during normal running, confirming that it is responsible for the propagation of leg swing.

Adult↗
PubMed2005

Independent metabolic costs of supporting body weight and accelerating body mass during walking.

The metabolic cost of walking is determined by many mechanical tasks, but the individual contribution of each task remains unclear. We hypothesized that the force generated to support body weight and the work performed to redirect and accelerate body mass each individually incur a significant metabolic cost during normal walking. To test our hypothesis, we measured changes in metabolic rate in response to combinations of simulated reduced gravity and added loading. We found that reducing body weight by simulating reduced gravity modestly decreased net metabolic rate. By calculating the metabolic cost per Newton of reduced body weight, we deduced that generating force to support body weight comprises approximately 28% of the metabolic cost of normal walking. Similar to previous loading studies, we found that adding both weight and mass increased net metabolic rate in more than direct proportion to load. However, when we added mass alone by using a combination of simulated reduced gravity and added load, net metabolic rate increased about one-half as much as when we added both weight and mass. By calculating the cost per kilogram of added mass, we deduced that the work performed on the center of mass comprises approximately 45% of the metabolic cost of normal walking. Our findings support the hypothesis that force and work each incur a significant metabolic cost. Specifically, the cost of performing work to redirect and accelerate the center of mass is almost twice as great as the cost of generating force to support body weight.

Acceleration↗
PubMed2005

Energy cost and muscular activity required for leg swing during walking.

To investigate the metabolic cost and muscular actions required for the initiation and propagation of leg swing, we applied a novel combination of external forces to subjects walking on a treadmill. We applied a forward pulling force at each foot to assist leg swing, a constant forward pulling force at the waist to provide center of mass propulsion, and a combination of these foot and waist forces to evaluate leg swing. When the metabolic cost and muscle actions were at a minimum, the condition was considered optimal. We reasoned that the difference in energy consumption between the optimal combined waist and foot force trial and the optimal waist force-only trial would reflect the metabolic cost of initiating and propagating leg swing during normal walking. We also reasoned that a lower muscle activity with these assisting forces would indicate which muscles are normally responsible for initiating and propagating leg swing. With a propulsive force at the waist of 10% body weight (BW), the net metabolic cost of walking decreased to 58% of normal walking. With the optimal combination, a propulsive force at the waist of 10% BW plus a pulling force at the feet of 3% BW the net metabolic cost of walking further decreased to 48% of normal walking. With the same combination, the muscle activity of the iliopsoas and rectus femoris muscles during the swing phase was 27 and 60% lower, respectively, but the activity of the medial gastrocnemius and soleus before swing did not change. Thus our data indicate that approximately 10% of the net metabolic cost of walking is required to initiate and propagate leg swing. Additionally, the hip flexor muscles contribute to the initiation and propagation leg swing.

Electromyography↗
PubMed2005

Giant Galapagos tortoises walk without inverted pendulum mechanical-energy exchange.

Animals must perform mechanical work during walking, but most conserve substantial mechanical energy via an inverted-pendulum-like mechanism of energy recovery in which fluctuations of kinetic energy (KE) and gravitational potential energy (GPE) are of similar magnitude and 180 degrees out of phase. The greatest energy recovery typically occurs at intermediate speeds. Tortoises are known for their slow speeds, which we anticipated would lead to small fluctuations in KE. To have an effective exchange of mechanical energy using the inverted-pendulum mechanism, tortoises would need to walk with only small changes in GPE corresponding to vertical center-of-mass (COM) fluctuations of < 0.5 mm. Thus, we hypothesized that giant Galapagos tortoises would not conserve substantial mechanical energy using the inverted-pendulum mechanism. We studied five adult giant Galapagos tortoises Geochelone elephantopus (mean mass=142 kg; range= 103-196 kg). Walking speed was extremely slow (0.16+/-0.052 m s(-1); mean +/- 1 s.d.). The fluctuations in kinetic energy (8.1+/-3.98 J stride(-1)) were only one-third as large as the fluctuations in gravitational potential energy (22.7+/-8.04 J stride(-1)). In addition, these energies fluctuated nearly randomly and were only sporadically out of phase. Because of the dissimilar amplitudes and inconsistent phase relationships of these energies, tortoises conserved little mechanical energy during steady walking, recovering only 29.8+/-3.77% of the mechanical energy (range=13-52%). Thus, giant Galapagos tortoises do not utilize effectively an inverted-pendulum mechanism of energy conservation. Nonetheless, the mass-specific external mechanical work required per distance (0.41+/-0.092 J kg(-1) m(-1)) was not different from most other legged animals. Other turtle species use less than half as much metabolic energy to walk as other terrestrial animals of similar mass. It is not yet known if Galapagos tortoises are economical walkers. Nevertheless, contrary to biomechanical convention, poor inverted-pendulum mechanics during walking do not necessarily correspond to high mechanical work and may not result in a high metabolic cost.

Animals↗
PubMed2004

Mechanical and metabolic requirements for active lateral stabilization in human walking.

Walking appears to be passively unstable in the lateral direction, requiring active feedback control for stability. The central nervous system may control stability by adjusting medio-lateral foot placement, but potentially with a metabolic cost. This cost increases with narrow steps and may affect the preferred step width. We hypothesized that external stabilization of the body would reduce the active control needed, thereby decreasing metabolic cost and preferred step width. To test these hypotheses, we provided external lateral stabilization, using springs pulling bilaterally from the waist, to human subjects walking on a force treadmill at 1.25 m/s. Ten subjects walked, with and without stabilization, at a prescribed step width of zero and also at their preferred step width. We measured metabolic cost using indirect calorimetry, and step width from force treadmill data. We found that at the prescribed zero step width, external stabilization resulted in a 33% decrease in step width variability (root-mean-square) and a 9.2% decrease in metabolic cost. In the preferred step width conditions, external stabilization caused subjects to prefer a 47% narrower step width, with a 32% decrease in step width variability and a 5.7% decrease in metabolic cost. These results suggest that (a). human walking requires active lateral stabilization, (b). body lateral motion is partially stabilized via medio-lateral foot placement, (c). active stabilization exacts a modest metabolic cost, and (d). humans avoid narrow step widths because they are less stable.

Adult↗
PubMed2004

Biomechanical and energetic determinants of the walk-trot transition in horses.

We studied nine adult horses spanning an eightfold range in body mass (M(b)) (90-720 kg) and a twofold range in leg length (L) (0.7-1.4 m). We measured the horses' walk-trot transition speeds using step-wise speed increments as they locomoted on a motorized treadmill. We then measured their rates of oxygen consumption over a wide range of walking and trotting speeds. We interpreted the transition speed results using a simple inverted-pendulum model of walking in which gravity provides the centripetal force necessary to keep the leg in contact with the ground. By studying a large size range of horses, we were naturally able to vary the absolute walking speed that would produce the same ratio of centripetal to gravitational forces. This ratio, (M(b)v2/L)/(M(b)g), reduces to the dimensionless Froude number (v2/gL), where v is forward speed, L is leg length and g is gravitational acceleration. We found that the absolute walk-trot transition speed increased with size from 1.6 to 2.3 m s(-1), but it occurred at nearly the same Froude number (0.35). In addition, horses spontaneously switched between gaits in a narrow range of speeds that corresponded to the metabolically optimal transition speed. These results support the hypotheses that the walk-trot transition is triggered by inverted-pendulum dynamics and occurs at the speed that maximizes metabolic economy.

Analysis of Variance↗
PubMed2003

Metabolic cost of generating muscular force in human walking: insights from load-carrying and speed experiments.

We sought to understand how leg muscle function determines the metabolic cost of walking. We first indirectly assessed the metabolic cost of swinging the legs and then examined the cost of generating muscular force during the stance phase. Four men and four women walked at 0.5, 1.0, 1.5, and 2.0 m/s carrying loads equal to 0, 10, 20, and 30% body mass positioned symmetrically about the waist. The net metabolic rate increased in nearly direct proportion to the external mechanical power during moderate-speed (0.5-1.5 m/s) load carrying, suggesting that the cost of swinging the legs is relatively small. The active muscle volume required to generate force on the ground and the rate of generating this force accounted for >85% of the increase in net metabolic rate across moderate speeds and most loading conditions. Although these factors explained less of the increase in metabolic rate between 1.5 and 2.0 m/s ( approximately 50%), the cost of generating force per unit volume of active muscle [i.e., the cost coefficient (k)] was similar across all conditions [k = 0.11 +/- 0.03 (SD) J/cm3]. These data indicate that, regardless of the work muscles do, the metabolic cost of walking can be largely explained by the cost of generating muscular force during the stance phase.

Adult↗
PubMed2002

Energy cost and muscular activity required for propulsion during walking.

We reasoned that with an optimal aiding horizontal force, the reduction in metabolic rate would reflect the cost of generating propulsive forces during normal walking. Furthermore, the reductions in ankle extensor electromyographic (EMG) activity would indicate the propulsive muscle actions. We applied horizontal forces at the waist, ranging from 15% body weight aiding to 15% body weight impeding, while subjects walked at 1.25 m/s. With an aiding horizontal force of 10% body weight, 1) the net metabolic cost of walking decreased to a minimum of 53% of normal walking, 2) the mean EMG of the medial gastrocnemius (MG) during the propulsive phase decreased to 59% of the normal walking magnitude, and yet 3) the mean EMG of the soleus (Sol) did not decrease significantly. Our data indicate that generating horizontal propulsive forces constitutes nearly half of the metabolic cost of normal walking. Additionally, it appears that the MG plays an important role in forward propulsion, whereas the Sol does not.

Adult↗
PubMed2002

Simultaneous positive and negative external mechanical work in human walking.

In human walking, the center of mass motion is similar to an inverted pendulum. Viewing double support as a transition from one inverted pendulum to the next, we hypothesized that the leading leg performs negative work to redirect the center of mass velocity, while simultaneously, the trailing leg performs positive work to replace the lost energy. To test this hypothesis, we developed a method to quantify the external mechanical work performed by each limb (individual limbs method). Traditional measures of external mechanical work use the sum of the ground reaction forces acting on the limbs (combined limbs method) allowing for the mathematical cancellation of simultaneous positive and negative work during multiple support periods. We expected to find that the traditional combined limbs method underestimates external mechanical work by a substantial amount. We used both methods to measure the external mechanical work performed by humans walking over a range of speeds. We found that during double support, the legs perform a substantial amount of positive and negative external work simultaneously. The combined limbs measures of positive and negative external work were approximately 33% less than those calculated using the individual limbs method. At all speeds, the trailing leg performs greater than 97% of the double support positive work while the leading leg performs greater than 94% of the double support negative work.

Biomechanical Phenomena↗
PubMed2002

Mechanical work for step-to-step transitions is a major determinant of the metabolic cost of human walking.

In the single stance phase of walking, center of mass motion resembles that of an inverted pendulum. Theoretically, mechanical work is not necessary for producing the pendular motion, but work is needed to redirect the center of mass velocity from one pendular arc to the next during the transition between steps. A collision model predicts a rate of negative work proportional to the fourth power of step length. Positive work is required to restore the energy lost, potentially exacting a proportional metabolic cost. We tested these predictions with humans (N=9) walking over a range of step lengths (0.4-1.1 m) while keeping step frequency fixed at 1.8 Hz. We measured individual limb external mechanical work using force plates, and metabolic rate using indirect calorimetry. As predicted, average negative and positive external mechanical work rates increased with the fourth power of step length (from 1 W to 38 W; r(2)=0.96). Metabolic rate also increased with the fourth power of step length (from 7 W to 379 W; r(2)=0.95), and linearly with mechanical work rate. Mechanical work for step-to-step transitions, rather than pendular motion itself, appears to be a major determinant of the metabolic cost of walking.

Biomechanical Phenomena↗