First published online March 28, 2008
Journal of Experimental Biology 211, 1211-1220 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.013227
Task-dependent force sharing between muscle synergists during locomotion in turkeys
Frank E. Nelson* and
Thomas J. Roberts
Ecology and Evolutionary Biology Department, Brown University, Box GB205,
Providence, RI 02912, USA

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Fig. 1. Swing duration for unloaded (open circles) running and for running with
added limb weights of 30 g (solid gray circles) and 60 g (solid black
circles). Values significantly different (P<0.05) from the
unloaded value at each speed are indicated with an asterisk. The symbols for
swing duration with limb weights of 30 g and 60 g are offset from their
measured speed to the right and left, respectively.
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Fig. 2. Values of peak force, time to peak force and impulse produced by the
lateral head of the gastrocnemius (red bars) during swing phase were not
different from the values required to generate the extension moment at the
ankle joint (gray bars) across speed and weighting conditions. The force and
impulse required did not change with added mass (P>0.09) and were
not significantly different (P>0.31) than the force produced by
the LG. For speeds 1–2 m s–1, Ni=5
for all variables; for 2.5 m s–1, Ni=4
for all variables.
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Fig. 3. A representative trace of joint moments (A) and forces (B,C) developed by
the MG (blue lines) and LG (red lines) for the right limb of a turkey running
at 2 m s–1. A positive then negative joint moment is required
at the ankle joint during swing as the joint flexes and extends. The black
line (B,C) is the force required from the gastrocnemius, calculated from the
moment measured by inverse dynamics and the muscle moment arm. Only forces
during swing phase were calculated from inverse dymanics (required force). The
muscular force required to produce the measured joint extension moment is
closely matched by the force produced by the lateral head of the
gastrocnemius, while the medial head produces almost no force. The exclusive
force production by the LG ends at the beginning of stance (gray shaded region
in C).
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Fig. 4. An equal amount of force is produced in the two heads of the gastrocnemius
during stance. Peak force significantly (P<0.05) increases in both
the lateral head (red) and the medial head (blue) with speed. With increasing
speed, the time of peak force occurs consistently later in the medial head.
Other timing variables were unchanged across changes in mass and speed. For
speeds 1–2 m s–1, Ni=5 for LG and
Ni=4 for MG variables. For 2.5 m s–1,
Ni=4 for LG variables and Ni=3 for MG
variables.
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Fig. 5. Reduced major axis regressions of MG peak force against LG peak forces
shows that for all speed and loading conditions, forces in the LG and MG are
similar during stance phase (A), but not during swing phase (B). Black
triangles, bird 1; red triangles, bird 2; green triangles, bird 3. The slopes
of both regression lines through the pooled data of all three birds were
significant (P<0.05). Solid lines, slopes for measured data;
broken line, hypothetical slope=1.0.
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© The Company of Biologists Ltd 2008