First published online December 28, 2007
Journal of Experimental Biology 211, 170-179 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.007476
Contractile properties of the pigeon supracoracoideus during different modes of flight
Bret W. Tobalske1,* and
Andrew A. Biewener2
1 Department of Biology, University of Portland, 5000 N. Willamette Boulevard,
Portland, OR 97203, USA
2 Concord Field Station, Harvard University, Old Causeway Road, Bedford, MA
01730, USA

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Fig. 1. Placement of (A) sonomicrometry crystals for measuring fascicle length in
the the supracoracoideus (SUPRA) and the pectoralis (PECT), and (B) strain
gauges on the dorsal surface of the deltopectoral crest of the left humerus
for measuring bone strain, calibrated to estimate SUPRA and PECT force, in a
pigeon (Columbia livia).
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Fig. 2. (A) Electromyographic (EMG) and contractile activity in the SUPRA and PECT
of a pigeon (Columbia livia) engaged in ascending flight (2.7 s);
standing on a platform (0–0.4 s), take-off and ascent to a perch
(0.4–2.0 s), and landing and resting on the perch (2.0–2.7 s). The
shaded area over fourth wingbeat highlights a region analyzed as representing
ascending flight. (B) Expanded view of data obtained during an ascending
wingbeat, corresponding to the shaded area in A.
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Fig. 3. Relative timing of length change, activation and force in the SUPRA and
PECT of flying pigeons (Columba livia, N=7). Data from different
modes of flight were pooled to create this figure. Values are means ±
s.d. Broken lines indicate data for a subsequent wingbeat.
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Fig. 4. Strain gauge recordings from the SUPRA and PECT of a pigeon (Columba
livia) illustrating antagonistic force development at the end of
downstroke and the end of upstroke. Overlap of force production was apparent
even when SUPRA recordings were not corrected for cross-talk from the
PECT.
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Fig. 5. (A) Peak stress ( m) in the SUPRA and PECT of pigeon
(Columba livia) during different modes of flight. (B) Fractional
length changes in the SUPRA and PECT according to mode of flight. Resting
length is indicated by the origin. Values are means ± s.d.,
N=7 (except ascending N=6).
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Fig. 6. Representative work loops from the SUPRA and PECT muscles of a pigeon
(Columba livia) engaged in (A,B) level, (C,D) ascending and (E,F)
descending flight. Arrows indicate direction of contraction; bold lines
indicate EMG activity. The hatched areas feature artificial negative stress
( m) due to compression of the SUPRA strain gauge by
cross-talk from PECT force that remained even after a correction factor was
applied; these areas were not included in the analysis.
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Fig. 7. Mean muscle power output (Pmus) measured in the SUPRA
and PECT muscles of the pigeon (Columba livia) during different modes
of flight compared with the estimated inertial power requirement
(Piner) for upstroke and aerodynamic power requirement
(Paero) assuming that all lift is provided during
downstroke. Values for Pmus are doubled to represent
output from paired left and right muscles. Values are means ± s.d.,
N=7 (except ascending N=6).
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© The Company of Biologists Ltd 2008