External work and potential for elastic storage at the limb joints of running dogs.

The storage and recovery of elastic strain energy in muscles and tendons increases the economy of locomotion in running vertebrates. In this investigation, we compared the negative and positive external work produced at individual limb joints of running dogs to evaluate which muscle-tendon systems contribute to elastic storage and to determine the extent to which the external work of locomotion is produced by muscles that shorten actively rather than by muscles that function as springs. We found that the negative and positive external work of the extensor muscles is not allocated equally among the different joints and limbs. During both trotting and galloping, the vast majority of the negative work was produced by the two distal joints, the wrist and ankle. The forelimb produced most of the negative work in both the trot and the gallop. The hindlimb produced most of the positive work during galloping, but not during trotting. With regards to elastic storage, our results indicate that the forelimb of dogs displays a greater potential for storage and recovery of elastic energy than does the hindlimb. Elastic storage appears to be more important during trotting than during galloping, and elastic storage appears to be more pronounced in the extensor muscles of the distal joints than in the extensor muscles of the proximal joints. Furthermore, our analysis indicates that a significant portion of the external work of locomotion, 26 % during trotting and 56 % during galloping, is produced by actively shortening muscles. We conclude that, although elastic storage of energy is extremely important to the economy of running gaits, actively shortening muscles do make an important contribution to the work of locomotion.

[1]  B. Katz The relation between force and speed in muscular contraction , 1939, The Journal of physiology.

[2]  G. Cavagna,et al.  MECHANICAL WORK IN RUNNING. , 1964, Journal of applied physiology.

[3]  G. Cavagna,et al.  The mechanics of sprint running , 1971, The Journal of physiology.

[4]  M. Hildebrand Analysis of Vertebrate Structure , 1974 .

[5]  G. Cavagna Force platforms as ergometers. , 1975, Journal of applied physiology.

[6]  B. Betts,et al.  Telemetered EMG of fast and slow muscles in cats , 1976, Brain Research.

[7]  G. Cavagna,et al.  Mechanical work in terrestrial locomotion: two basic mechanisms for minimizing energy expenditure. , 1977, The American journal of physiology.

[8]  R. Alexander,et al.  Mechanics of locomotion of dogs (Canis familiaris) and sheep (Ovis aries). , 2009, Journal of zoology.

[9]  G. Cavagna,et al.  Energetics and mechanics of terrestrial locomotion. III. Energy changes of the centre of mass as a function of speed and body size in birds and mammals. , 1982, The Journal of experimental biology.

[10]  N. Heglund,et al.  Energetics and mechanics of terrestrial locomotion. , 1982, Annual review of physiology.

[11]  N. Heglund,et al.  Energetics and mechanics of terrestrial locomotion. I. Metabolic energy consumption as a function of speed and body size in birds and mammals. , 1982, The Journal of experimental biology.

[12]  G. Cavagna,et al.  Energetics and mechanics of terrestrial locomotion. IV. Total mechanical energy changes as a function of speed and body size in birds and mammals. , 1982, The Journal of experimental biology.

[13]  R. M. Alexander Elastic Energy Stores in Running Vertebrates , 1984 .

[14]  M. Bobbert,et al.  An estimation of power output and work done by the human triceps surae muscle-tendon complex in jumping. , 1986, Journal of biomechanics.

[15]  M. Bobbert,et al.  A model of the human triceps surae muscle-tendon complex applied to jumping. , 1986, Journal of biomechanics.

[16]  R. F. Ker,et al.  The spring in the arch of the human foot , 1987, Nature.

[17]  R. M. Alexander,et al.  Elastic mechanisms in animal movement , 1988 .

[18]  R. Blickhan The spring-mass model for running and hopping. , 1989, Journal of biomechanics.

[19]  M. Bobbert,et al.  The Unique Action of Bi-Articular Muscles in Leg Extensions , 1990 .

[20]  T. McMahon,et al.  The mechanics of running: how does stiffness couple with speed? , 1990, Journal of biomechanics.

[21]  R. Alexander,et al.  A model of bipedal locomotion on compliant legs. , 1992, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[22]  J. Faulkner,et al.  Injury to skeletal muscle fibers during contractions: conditions of occurrence and prevention. , 1993, Physical therapy.

[23]  C. T. Farley,et al.  Running springs: speed and animal size. , 1993, The Journal of experimental biology.

[24]  C. R. Taylor,et al.  Relating mechanics and energetics during exercise. , 1994, Advances in veterinary science and comparative medicine.

[25]  T. Cr Relating mechanics and energetics during exercise. , 1994 .

[26]  V M Zatsiorsky,et al.  Tendon action of two-joint muscles: transfer of mechanical energy between joints during jumping, landing, and running. , 1994, Journal of biomechanics.

[27]  D. Carrier Function of the intercostal muscles in trotting dogs: ventilation or locomotion? , 1996, The Journal of experimental biology.

[28]  W Herzog,et al.  Transfer of mechanical energy between ankle and knee joints by gastrocnemius and plantaris muscles during cat locomotion. , 1996, Journal of biomechanics.

[29]  T J Roberts,et al.  Muscular Force in Running Turkeys: The Economy of Minimizing Work , 1997, Science.

[30]  D R Carrier,et al.  Dynamic gearing in running dogs. , 1998, The Journal of experimental biology.

[31]  R L Marsh,et al.  Activation patterns and length changes in hindlimb muscles of the bullfrog Rana catesbeiana during jumping. , 1998, The Journal of experimental biology.