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First published online June 6, 2005
Journal of Experimental Biology 208, 2377-2387 (2005)
Published by The Company of Biologists 2005
doi: 10.1242/jeb.01618
Patterns of red muscle strain/activation and body kinematics during steady swimming in a lamnid shark, the shortfin mako (Isurus oxyrinchus)
1 Marine Biology Research Division, Scripps Institution of Oceanography,
University of California, San Diego, La Jolla, CA 92093-0202, USA
2 Pfleger Institute of Environmental Research, Oceanside, CA 92054,
USA
3 Department of Zoology, University of Tübingen, Auf der Morgenstelle
28, 72076 Tübingen, Germany
* Author for correspondence (e-mail: jdonley{at}ucsd.edu)
Accepted 29 March 2005
The dynamics of steady swimming were examined in the shortfin mako
(Isurus oxyrinchus), a member of the cartilaginous fish family
Lamnidae, a family known for their morphological adaptations for
high-performance locomotion and their similarity in hydromechanical design to
tunas. Patterns of red muscle (RM) strain (i.e. relative length change) and
activation were quantified at two axial positions (
0.4 and 0.6L,
where L is total body length), using sonomicrometry and
electromyography (EMG), and correlated with simultaneous measurements of
dorsal midline kinematics during steady swimming (
0.51 L
s1). RM strain varied longitudinally with strain amplitudes
ranging from 5.5±1.1% (S.E.M.) in the anterior to
8.7±0.9% in the posterior. We found no significant longitudinal
variation in patterns of RM activation, with mean onset of activation
occurring at 8384° (90° is peak length) and offset at
200210° at both body positions. Likewise, duty cycles were similar:
35.5±1.0% in the anterior and 32.2±1.6% in the posterior.
Comparison of the timing of waves of dorsal midline curvature and predicted
strain relative to measured RM strain revealed a phase shift between RM
shortening and local body bending. Furthermore, when the body is bent
passively, RM shortens synchronously with the surrounding white muscle (WM)
and skin, as expected. During active swimming, peaks in RM strain were delayed
relative to peaks in WM strain by a mean of
10% of the tailbeat cycle,
with one individual as high as
17% in the anterior and nearly 50% in the
posterior. The longitudinal consistency in the EMG/strain phase relationship
in the mako is similar to that in the leopard shark, suggesting a consistent
trend among sharks using different locomotor modes. However, unlike in the
leopard shark, RM shortening in the mako is physically uncoupled from
deformation of the surrounding body during steady swimming, a characteristic
shared between the mako and tunas.
Key words: muscle activation, strain, swimming, lamnid, Isurus
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