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First published online December 14, 2007
Journal of Experimental Biology 211, 150-162 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.010678
Locomotor function of forelimb protractor and retractor muscles of dogs: evidence of strut-like behavior at the shoulder
1 Department of Biology, 201 South Biology Building, University of Utah, Salt
Lake City, UT 84112, USA
2 Department of Biology, 4202 East Fowler Avenue, SCA 110, University of South
Florida, Tampa, FL 33620, USA
* Author for correspondence (e-mail: carrier{at}biology.utah.edu)
Accepted 18 October 2007
The limbs of running mammals are thought to function as inverted struts. When mammals run at constant speed, the ground reaction force vector appears to be directed near the point of rotation of the limb on the body such that there is little or no moment at the joint. If this is true, little or no external work is done at the proximal joints during constant-speed running. This possibility has important implications to the energetics of running and to the coupling of lung ventilation to the locomotor cycle. To test if the forelimb functions as an inverted strut at the shoulder during constant-speed running and to characterize the locomotor function of extrinsic muscles of the forelimb, we monitored changes in the recruitment of six muscles that span the shoulder (the m. pectoralis superficialis descendens, m. pectoralis profundus, m. latissimus dorsi, m. omotransversarius, m. cleidobrachialis and m. trapezius) to controlled manipulations of locomotor forces and moments in trotting dogs (Canis lupus familiaris Linnaeus 1753). Muscle activity was monitored while the dogs trotted at moderate speed (approximately 2 m s–1) on a motorized treadmill. Locomotor forces were modified by (1) adding mass to the trunk, (2) inclining the treadmill so that the dogs ran up- and downhill (3) adding mass to the wrists or (4) applying horizontally directed force to the trunk through a leash. When the dogs trotted at constant speed on a level treadmill, the primary protractor muscles of the forelimb exhibited activity during the last part of the ipsilateral support phase and the beginning of swing phase, a pattern that is consistent with the initiation of swing phase but not with active protraction of the limb during the beginning of support phase. Results of the force manipulations were also consistent with the protractor muscles initiating swing phase and contributing to active braking via production of a protractor moment on the forelimb when the dogs decelerate. A similar situation appears to be true for the major retractor muscles of the forelimb. The m. pectoralis profundus and the m. latissimus dorsi were completely silent during the support phase of the ipsilateral limb when the dogs ran unencumbered and exhibited little or no increase in activity when the dogs carried added mass on their backs to increase any retraction torque during the support phase of constant-speed running. The most likely explanation for these observations is that the ground force reaction vector is oriented very close to the fulcrum of the forelimb such that the forelimb functions as a compliant strut at the shoulder when dogs trot at constant speed on level surfaces. Because the moments at the fulcrum of the pectoral girdle appear to be small during the support phase of a trotting step, a case can be made that it is the activity of the extrinsic appendicular muscles that produce the swing phase of the forelimb that explain the coupled phase relationship between ventilatory airflow and the locomotor cycle in trotting dogs.
Key words: forelimb, retractor muscles, protractor muscles, EMG, recruitment, quadruped, running, shoulder
