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First published online July 17, 2009
Journal of Experimental Biology 212, 2454-2463 (2009)
Published by The Company of Biologists 2009
doi: 10.1242/jeb.027995
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Intensity of activation and timing of deactivation modulate elastic energy storage and release in a pennate muscle and account for gait-specific initiation of limb protraction in the horse

Glen A. Lichtwark1,*, Johanna C. Watson1, Sophia Mavrommatis2 and Alan M. Wilson1

1 Structure and Motion Laboratory, The Royal Veterinary College, Hawkshead Lane, North Mymms, Hatfield, Hertfordshire AL9 7TA, UK
2 Royal National Orthopaedic Hospital, Brockley Hill, Stanmore, Middlesex HA7 4LP, UK

* Author for correspondence at present address: School of Physiotherapy and Exercise Science, Griffith University, Gold Coast, Queensland 4222, Australia (e-mail: g.lichtwark{at}griffith.edu.au)

Accepted 1 May 2009

The equine biceps brachii (biceps) initiates rapid limb protraction through a catapult mechanism. Elastic strain energy is slowly stored in an internal tendon and is then rapidly released to protract the forelimb. The muscle fibres are short, have little scope for length change and can therefore only shorten slowly compared with the speed at which the whole muscle must shorten, which makes them poor candidates for driving rapid limb protraction. We suggest that the muscle fibres in the biceps act to modulate the elastic energy output of the muscle–tendon unit (MTU) to meet the demands of locomotion under different conditions. We hypothesise that more elastic strain energy is stored and released from the biceps MTU during higher speed locomotion to accommodate the increase in energy required to protract the limb and that this can be achieved by varying the length change and activation conditions of the muscle. We examined the work performed by the biceps during trot and canter using an inverse dynamics analysis (IDA). We then used excised biceps muscles to determine how much work could be performed by the muscle in active and passive stretch–shorten cycles. A muscle model was developed to investigate the influence of changes in activation parameters on energy storage and energy return from the biceps MTU. Increased biceps MTU length change and increased work performed by the biceps MTU were found at canter compared with at trot. More work was performed by the ex vivo biceps MTU following activation of the muscle and by increasing muscle length change. However, the ratio of active to passive work diminished with increasing length change. The muscle model demonstrated that duration and timing of activation during stretch–shorten cycles could modulate the elastic energy storage and return from the biceps. We conclude that the equine biceps MTU acts as a tuneable spring and the contractile component functions to modulate the energy required for rapid forelimb protraction at different speeds.

Key words: muscle, elasticity, locomotion, biomechanics, horse


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© The Company of Biologists Ltd 2009