First published online August 8, 2008
Journal of Experimental Biology 211, 2669-2677 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.015883
The `upstream wake' of swimming and flying animals and its correlation with propulsive efficiency
Jifeng Peng1,* and
John O. Dabiri1,2
1 Bioengineering, California Institute of Technology, Pasadena, CA 91125,
USA
2 Graduate Aeronautical Laboratories, California Institute of Technology,
Pasadena, CA 91125, USA

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Fig. 1. Schematic flexible plate model swimmer. s is the curve length and
(t,s) is the tangential angle. LE, leading edge
(s=–1); TE, trailing edge (s=1).
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Fig. 3. Schematic of particle separation at the boundary of a vortex ring. (A)
Fluid particle pairs straddling the vortex boundary have a larger separation
rate in backward time, indicating larger values of backward-time FTLE
(finite-time Lyapunov exponent) at the front boundary of the vortex ring. (B)
Fluid particle pairs straddling the vortex boundary have a larger separation
rate in forward time, indicating larger values of forward-time FTLE at the
rear boundary of the vortex ring. x and y are particle
trajectories; T is integration time.
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Fig. 4. The effect of integration time on FTLE (finite-time Lyapunov exponent) and
LCS (Lagrangian Coherent Structure) calculation for a vortex ring. (A)
integration time T=0.4 s; (B) T=1.2 s; (C) T=2.0 s;
(D) T=2.8 s. With longer integration time, the FTLE ridges become
sharper, i.e. LCS resolve into clearly defined thin lines.
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Fig. 5. Locomotion of the model swimmer. The swimmer (S) begins at rest (A) and
flaps its flexible body to propel itself forward. The vortex wake generated by
the swimming motion (blue curve) and the resulting forward motion of the
swimmer are shown after two (B), four (C) and six (D) swimming cycles. See
Movie 2 in supplementary material.
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Fig. 6. Finite-time Lyapunov exponent (FTLE) fields of the flow created by the
model swimmer (S). Results are presented in a reference frame fixed on the
swimmer; the swimmer moves from right to left in a laboratory reference frame
(see Fig. 5). Top panel, time
t=0; second panel, t=T/5; third panel,
t=(2/5)T; bottom panel, t=(4/5)T, where
T is the duration of a single swimming cycle. (A) Backward-time FTLE
field. The ridge of large FTLE values (solid blue curve) identifies the
attracting LCS. (B) Forward-time FTLE field. The ridge of large FTLE values
(solid red curve) identifies the repelling LCS. See Movie 3 in supplementary
material.
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Fig. 7. Temporal evolution of the upstream wake. (A) Time t=0; (B)
t=(7/5)T; (C) t =(14/5)T, where T
is the duration of a single swimming cycle. See Movie 4 in supplementary
material. Fluid particles in each of three adjacent repelling LCS (red curves)
are labeled magenta or green in order to track their evolution. After
interaction with the swimmer (shown here in a reference frame fixed on the
swimmer; the swimmer moves from right to left in a laboratory reference frame;
see Fig. 5), the fluid
particles in the repelling LCS are shown to comprise the subsequent downstream
wake, illustrated by the attracting LCS (blue curves). Fluid particles
initially separated by the repelling LCS do not mix and are only moderately
deformed. S, swimmer; w(x), the width of the upstream wake; RLCS,
repelling LCS; ALCS, attracting LCS.
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Fig. 8. Temporal evolution of arbitrary upstream fluid parcels. Fluid particles in
adjacent regions with a horizontal (A,B) or vertical (C,D) interface are
labeled magenta or green in order to track their evolution. The interfaces in
both cases do not coincide with the repelling LCS (red curves). After
interaction with the swimmer (shown here in a reference frame fixed on the
swimmer; the swimmer moves from right to left in a laboratory reference frame;
see Fig. 5), the particles in
the adjacent parcels exhibit substantial deformation and mixing in the
vicinity of the attracting LCS (blue curves). In both cases, the parcels do
not indicate the full extent of the downstream wake, in contrast to the
repelling LCS parcels in Fig.
7. S, swimmer; RLCS, repelling LCS; ALCS, attracting LCS. (A,C)
Time t=0; (B,D) t=(14/5)T, where T is the
duration of a single swimming cycle. See Movies 5 and 6 in supplementary
material.
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Fig. 9. Total force acting on the swimmer (A) and the velocity of the swimmer (B).
The swimmer starts from rest. As it approaches a steady state, i.e. the mean
velocity over a stroke cycle approaches a constant, the time average of force
approaches zero.
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Fig. 10. Efficiency based on mass flow rate (Eqn
6) of the model swimmer versus Strouhal number
St=fA/ . Red, kinematics 1; black, kinematics
2; green, kinematics 3. Error bars indicate uncertainty in measurement of
upstream wake width.
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Fig. 11. Comparison of measures for swimming efficiency. Plots A, B and C show
kinematics 1, 2 and 3, respectively. Solid lines: efficiency based on the
width of the upstream fluid structure; error bars indicate uncertainty in
measurement of upstream wake width; broken lines: efficiency calculated by
replacing upstream wake width with stroke amplitude of the swimmer
(peak-to-peak excursion at trailing edge).
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Fig. 12. The width of the wake. (A) Wake width of a swimmer swimming at a constant
velocity (2 BL s–1) with different tail amplitudes. (B) Wake
width of a swimmer swimming at different velocities with the same body
kinematics (flapping amplitudes of 1.44 BL).
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Fig. 13. Efficiency of the swimmer versus modified Strouhal number
St=fw/ . Red, kinematics 1; black, kinematics
2; green, kinematics 3.
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© The Company of Biologists Ltd 2008