Effect of variation in form on the cost of terrestrial locomotion

1990 ◽  
Vol 150 (1) ◽  
pp. 233-246 ◽  
Author(s):  
R. J. Full ◽  
D. A. Zuccarello ◽  
A. Tullis

The mass-specific minimum cost of terrestrial locomotion (Cmin) decreases with an increase in body mass. This generalization spans nearly eight orders of magnitude in body mass and includes two phyla. The general relationship between metabolic cost and mass is striking. However, a significant amount of unexplained interspecific variation in Cmin exists at any given body mass. To determine how variation in morphology and physiology affects metabolic energy cost, we measured the oxygen consumption of three comparably sized insects running on a miniature treadmill; the American cockroach Periplaneta americana, the caterpillar hunting beetle Calosoma affine and the Australian field cricket Teleogryllus commodus. Steady-state oxygen consumption (VO2ss) increased linearly with speed. Cmin was similar for crickets and cockroaches (8.0 and 8.5 ml O2 g-1km-1, respectively), but was substantially lower for beetles (4.6 ml O2 g-1km-1). The predicted value of Cmin for all three insects was within the 95% confidence intervals of the Cmin versus body mass function. However, the 95% confidence intervals extend approximately 2.5-fold above and 40% below the regression line, making the variation at any given body mass nearly sixfold. Normalizing for the rate of muscle force production by determining the metabolic cost per stride failed to account for the interspecific variation in the cost of locomotion observed in the three insects. Ground contact costs (i.e. VO2ss multiplied by leg contact time during a stride) in insects were similar to those measured in mammals (1.5-3.1 J kg-1) and were independent of speed, but did not explain the interspecific variation in the cost of locomotion. Muscles of the caterpillar hunting beetle may have a greater mechanical advantage than muscles of the Australian field cricket and American cockroach. Variation in musculo-skeletal arrangement, apart from variation in body mass, could translate into significant differences in the minimum cost of terrestrial locomotion.

1990 ◽  
Vol 149 (1) ◽  
pp. 307-317 ◽  
Author(s):  
R. J. Full ◽  
A. Tullis

Small animals use more metabolic energy per unit mass than large animals to run on a level surface. If the cost to lift one gram of mass one vertical meter is constant, small animals should require proportionally smaller increases in metabolic cost to run uphill. To test this hypothesis on very small animals possessing an exceptional capacity for ascending steep gradients, we measured the metabolic cost of locomotion in the cockroach, Periplaneta americana, running at angles of 0, 45 and 90 degrees to the horizontal. Resting oxygen consumption (VO2rest) was not affected by incline angle. Steady-state oxygen consumption (VO2ss) increased linearly with speed at all angles of ascent. The minimum cost of locomotion (the slope of the VO2ss versus speed function) increased with increasing angle of ascent. The minimum cost of locomotion on 45 and 90 degrees inclines was two and three times greater, respectively, than the cost during horizontal running. The cockroach's metabolic cost of ascent greatly exceeds that predicted from the hypothesis of a constant efficiency for vertical work. Variations in stride frequency and contact time cannot account for the high metabolic cost, because they were independent of incline angle. An increase in the metabolic cost or amount of force production may best explain the increase in metabolic cost. Small animals, such as P. americana, can easily scale vertical surfaces, but the energetic cost is considerable.


1970 ◽  
Vol 53 (3) ◽  
pp. 763-777 ◽  
Author(s):  
HENRY D. PRANGE ◽  
KNUT SCHMIDT-NIELSEN

1. The metabolic cost of swimming was studied in mallard ducks (Anas platyrhynchos) which had been trained to swim steadily in a variable-speed water channel. 2. At speeds of from 0.35 to 0.50 m/sec the oxygen consumption remained relatively constant at approximately 2.2 times the resting level. At speeds of 0.55 m/sec and higher the oxygen consumption increased rapidly and reached 4.1 times resting at the maximum sustainable speed of 0.70 m/sec. 3. The maximum sustainable swimming speed of the ducks coincided with the limit predicted from hydrodynamic considerations of the water resistance of a displacement-hulled ship of the same hull length as a duck (0.33 m). 4. The cost of transport (metabolic rate/speed) reached a minimum of 5.77 kcal/kg km at a swimming speed of 0.50 m/sec. Ducks swimming freely on a pond were observed to swim at the speed calculated in experimental trials to give minimum cost of transport. 5. Drag measurements made with model ducks indicated a maximum overall efficiency (power output/power input) for the swimming ducks of about 5%. Ships typically have maximum efficiencies of 20-30%. Because of the difficulty in delimiting the cost of swimming activity alone from the other bodily functions of the duck, overall efficiency may present an incorrect description of the swimming performance of the duck relative to that of a ship. An hydrodynamic parameter such as speed/length ratio [speed/(hull length)½] whereby a duck excels conventional ships may present a more appropriate comparison.


1988 ◽  
Vol 138 (1) ◽  
pp. 301-318 ◽  
Author(s):  
N. C. Heglund ◽  
C. R. Taylor

In this study we investigate how speed and stride frequency change with body size. We use this information to define ‘equivalent speeds’ for animals of different size and to explore the factors underlying the six-fold difference in mass-specific energy cost of locomotion between mouse- and horse-sized animals at these speeds. Speeds and stride frequencies within a trot and a gallop were measured on a treadmill in 16 species of wild and domestic quadrupeds, ranging in body size from 30 g mice to 200 kg horses. We found that the minimum, preferred and maximum sustained speeds within a trot and a gallop all change in the same rather dramatic manner with body size, differing by nine-fold between mice and horses (i.e. all three speeds scale with about the 0.2 power of body mass). Although the absolute speeds differ greatly, the maximum sustainable speed was about 2.6-fold greater than the minimum within a trot, and 2.1-fold greater within a gallop. The frequencies used to sustain the equivalent speeds (with the exception of the minimum trotting speed) scale with about the same factor, the −0.15 power of body mass. Combining this speed and frequency data with previously published data on the energetic cost of locomotion, we find that the mass-specific energetic cost of locomotion is almost directly proportional to the stride frequency used to sustain a constant speed at all the equivalent speeds within a trot and a gallop, except for the minimum trotting speed (where it changes by a factor of two over the size range of animals studied). Thus the energy cost per kilogram per stride at five of the six equivalent speeds is about the same for all animals, independent of body size, but increases with speed: 5.0 J kg-1 stride-1 at the preferred trotting speed; 5.3 J kg-1 stride-1 at the trot-gallop transition speed; 7.5 J kg-1 stride-1 at the preferred galloping speed; and 9.4 J kg-1 stride-1 at the maximum sustained galloping speed. The cost of locomotion is determined primarily by the cost of activating muscles and of generating a unit of force for a unit of time. Our data show that both these costs increase directly with the stride frequency used at equivalent speeds by different-sized animals. The increase in cost per stride with muscles (necessitating higher muscle forces for the same ground reaction force) as stride length increases both in the trot and in the gallop.


1999 ◽  
Vol 86 (5) ◽  
pp. 1657-1662 ◽  
Author(s):  
Young-Hui Chang ◽  
Rodger Kram

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 (V˙o 2) 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,V˙o 2 increased 30% compared with normal running, presumably because of the extra work involved. With an AHF of +15% BWt, the subjects exerted ∼70% less propulsive impulse and exhibited a 33% reduction inV˙o 2. Our data suggest that generating horizontal propulsive forces constitutes more than one-third of the total metabolic cost of normal running.


1996 ◽  
Vol 199 (3) ◽  
pp. 587-592 ◽  
Author(s):  
C Farley ◽  
M Emshwiller

Nocturnal geckos can walk on level ground more economically than diurnal lizards. One hypothesis for why nocturnal geckos have a low cost of locomotion is that they can perform mechanical work during locomotion more efficiently than other lizards. To test this hypothesis, we compared the efficiency of the nocturnal gecko Coleonyx variegatus (average body mass 4.2 g) and the diurnal skink Eumeces skiltonianus (average body mass 4.8 g) when they performed vertical work during uphill locomotion. We measured the rate of oxygen consumption when each species walked on the level and up a 50 slope over a range of speeds. For Coleonyx variegatus, the energetic cost of traveling a unit distance (the minimum cost of transport, Cmin) increased from 1.5 to 2.7 ml O2 kg-1 m-1 between level and uphill locomotion. For Eumeces skiltonianus, Cmin increased from 2.5 to 4.7 ml O2 kg-1 m-1 between level and uphill locomotion. By taking the difference between Cmin for level and uphill locomotion, we found that the efficiency of performing vertical work during locomotion was 37 % for Coleonyx variegatus and 19 % for Eumeces skiltonianus. The similarity between the 1.9-fold difference in vertical efficiency and the 1.7-fold difference in the cost of transport on level ground is consistent with the hypothesis that nocturnal geckos have a lower cost of locomotion than other lizards because they can perform mechanical work during locomotion more efficiently.


1992 ◽  
Vol 262 (5) ◽  
pp. R771-R778 ◽  
Author(s):  
R. V. Baudinette ◽  
G. K. Snyder ◽  
P. B. Frappell

Rates of oxygen consumption and blood lactate levels were measured in tammar wallabies (Macropus eugenii) trained to hop on a treadmill. In addition, the work required to overcome wind resistance during forward locomotion was measured in a wind tunnel. Up to approximately 2.0 m/s, rates of oxygen consumption increased linearly with speed and were not significantly different from rates of oxygen consumption for a quadruped of similar body mass. Between 2.0 and 9.4 m/s, rates of oxygen consumption were independent of hopping speed, and between 3.9 and 7.9 m/s, the range over which samples were obtained, blood lactate levels were low (0.83 +/- 0.13 mmol.min-1.kg-1) and did not increase with hopping speed. The work necessary to overcome drag increased exponentially with speed but increased the energy cost of locomotion by only 10% at the average speed attained by our fast hoppers. Thus, during hopping, the energy cost of locomotion is effectively independent of speed. At rates of travel observed in the field, the estimated energy cost of transport in large macropods is less than one-third the cost for a quadruped of equivalent body mass. The energetic savings associated with this unique form of locomotion may have been an important physiological adaptation, enabling large macropods to efficiently cover the distances necessary to forage in the semiarid landscapes of Australia.


Physiology ◽  
1986 ◽  
Vol 1 (5) ◽  
pp. 153-155
Author(s):  
CR Taylor

African women can carry loads of 20% of their body mass on their heads without measurable metabolic cost. In contrast, trained or untrained humans and animals increase their metabolism by 20% when they carry loads of this magnitude. Two possible mechanisms are proposed for the women's economic means of load carrying, based on considerations of the cost of generating muscular force during locomotion.


2001 ◽  
Vol 204 (4) ◽  
pp. 797-803 ◽  
Author(s):  
F.E. Fish ◽  
P.B. Frappell ◽  
R.V. Baudinette ◽  
P.M. MacFarlane

The platypus Ornithorhynchus anatinus Shaw displays specializations in its limb structure for swimming that could negatively affect its terrestrial locomotion. Platypuses walked on a treadmill at speeds of 0.19-1.08 m × s(−1). Video recordings were used for gait analysis, and the metabolic rate of terrestrial locomotion was studied by measuring oxygen consumption. Platypuses used walking gaits (duty factor >0.50) with a sprawled stance. To limit any potential interference from the extensive webbing on the forefeet, platypuses walk on their knuckles. Metabolic rate increased linearly over a 2.4-fold range with increasing walking speed in a manner similar to that of terrestrial mammals, but was low as a result of the relatively low standard metabolic rate of this monotreme. The dimensionless cost of transport decreased with increasing speed to a minimum of 0.79. Compared with the cost of transport for swimming, the metabolic cost for terrestrial locomotion was 2.1 times greater. This difference suggests that the platypus may pay a price in terrestrial locomotion by being more aquatically adapted than other semi-aquatic or terrestrial mammals.


1993 ◽  
Vol 179 (1) ◽  
pp. 245-259 ◽  
Author(s):  
D. Berrigan ◽  
J. R. Lighton

We report the cost of transport and kinematics of terrestrial locomotion by larval blowflies (Protophormia terraenovae, Diptera: Calliphoridae). We contrast inter- and intra-individual methods for estimating minimum cost of transport (MCOT) and the relationship between speed, contraction frequency and distance traveled per contraction. The minimum cost of transport calculated from intra-individual data is 2297 +/− 317 J kg-1 m-1 (S.E.M.) and the MCOT calculated from inter-individual comparisons is statistically indistinguishable at 1910 +/− 327 J kg-1 m-1. These values are almost ten times higher than the predicted value for a similar-sized limbed arthropod. Fly larvae travel by repeated peristaltic contractions and individual contractions cost about the same amount as individual strides in limbed insects. Both contraction frequency and distance traveled per contraction increase linearly with speed. Doubling the contraction frequency or the distance traveled per contraction approximately doubles speed. The cost of transport in fly larvae is among the highest recorded for terrestrial locomotion, confirming the suggestion that biomechanical and kinematic properties of limbless organisms with hydraulic skeletons lead to very high costs of transport.


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