First published online November 14, 2008
Journal of Experimental Biology 211, 3661-3670 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.018754
The mechanics of the gibbon foot and its potential for elastic energy storage during bipedalism
Evie E. Vereecke1,2,* and
Peter Aerts2,3
1 Department of Human Anatomy and Cell Biology, School of Biomedical Sciences,
University of Liverpool, Liverpool L69 3GE, UK
2 Laboratorium for Functional Morphology, University of Antwerp,
Universiteitsplein 1, B-2610 Antwerp, Belgium
3 Department of Movement and Sports Sciences, University of Ghent,
Watersportlaan 2, B-9000 Gent, Belgium

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Fig. 1. Diagram of the gibbon foot indicating the major anatomical structures. Ga,
gastrocnemius (part of the triceps); FF+FT, flexor fibularis and flexor
tibialis (the long digital flexors); FDB, flexor digitorum brevis (the short
digital flexors); PL, plantar ligaments; AT, Achilles' tendon.
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Fig. 2. The four-linked segment foot model. (A) Video image of the gibbon foot with
indication of the digitized points. (B) Diagram of the joint angles. K, knee
joint; TC, talocrural joint; H, heel; TM, tarsometatarsal joint; MP,
metatarsophalangeal joint; T, toes.
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Fig. 4. Sagittal foot motion during the stance phase, demonstrating, from left to
right, touchdown, loading, heel-rise and push-off. Shown are three
representative bipedal bouts of three different individuals (top, young adult
male; middle, adult male; bottom, adult female – note heel-elevated
position during loading phase in the latter).
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Fig. 5. Stick figure of an average stance phase showing the four-segment link foot
model and the ground reaction force vector. (A) Touchdown (0–10% stance
phase), (B) loading phase (10–50% stance phase), (C) heel-rise
(50–80% stance phase), and (D) push-off (80–100% stance phase);
see text for details. [Note that the initial contact (0–10% stance
phase) is typically made with a widely abducted hallux, not with the hindfoot
(H–TM segment) although this illustration and
Fig. 6 do not give this
impression.]
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Fig. 6. Butterfly (Pedotti) diagram illustrating the ground reaction force vectors
as a function of foot length. The colours of the arrows indicate timing:
orange arrows, 0–10% stance phase; green arrows, 10–50%; blue
arrows, 50–80%; and red arrows, 80–100%. TM, tarsometatarsal
joint; MP, metatarsophalangeal joint. Arrows pointing to the left denote
braking, arrows pointing to the right denote acceleration.
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Fig. 7. Combination of average joint angles (white line; grey shaded area indicates
standard deviation) and moments at the TC (top, dashed line), TM (middle,
dotted line) and MP(bottom, solid line) joints against stance time (%).
Plantarflexor moments are positive, dorsiflexor moments negative; a decreasing
angle means dorsiflexion, an increase points to plantarflexion. Shaded areas
on the right denote positive power output (i.e. plantarflexor moment+joint
plantarflexion).
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Fig. 8. Instantaneous powers performed at the MP, TM and TC joints and the total
foot power (SUM) during stance. The bars at the bottom indicate foot contact
of the stance foot (black) and contralateral foot (grey).
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Fig. 9. Power performed at the foot (sum of MP, TM and TC joint power; light blue)
and total power performed to move the centre of mass (dark blue) during stance
with estimates of positive and negative external work.
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© The Company of Biologists Ltd 2008