First published online May 18, 2006
Journal of Experimental Biology 209, 2025-2033 (2006)
Published by The Company of Biologists 2006
doi: 10.1242/jeb.02242
Fast-swimming hydromedusae exploit velar kinematics to form an optimal vortex wake
John O. Dabiri1,*,
Sean P. Colin2 and
John H. Costello3
1 Graduate Aeronautical Laboratories and Bioengineering, California
Institute of Technology, Mail Code 138-78, Pasadena, CA 91125, USA
2 Biology and Marine Biology, Roger Williams University, MNS 241, Bristol,
RI 02809, USA
3 Biology, Providence College, Providence, RI 02918, USA

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Fig. 1. Morphological analysis of N. bachei. (A,B) Images of N.
bachei (A) at rest, and (B) during jet ejection. Image height is 7.2 mm.
V, velum; DV, velar diameter; T, tentacles; OC, oral cavity; MW,
mesogleal wall; A, animal apex. Out-pocketing of velum into funnel shape is
discernable in B. (C) Sample of image analysis. Image from B superimposed with
reconstructions of the oral cavity boundary (broken black line) and velum
(solid black line), based on control points (white circles).
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Fig. 2. Measurements of N. bachei body kinematics during fast swimming.
Two sets of measurements from distinct swimming cycles from two different
animals (black and grey circles) are presented to indicate the repeatability
of the swimming motions. Maximum measurement uncertainty is ±6%. A
curve fit to the data (solid black line) is used in subsequent analyses. (A)
Velar diameter versus time during the swimming cycle. (B) Oral cavity
volume versus time during the swimming cycle.
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Fig. 3. Plot of vortex formation time (T*) during the jet
ejection phase (t/Tejection) of the swimming
cycle (solid black line). At the end of the ejection phase (black star), the
vortex formation time coincides with the range of values producing maximum
vortex growth in mechanically generated jet flows with similar aperture
kinematics (grey band) (Dabiri and Gharib,
2005b ), where the aperture diameter contraction rate ranges
between 15% and 30% of the average jet velocity. For comparison, the measured
velar aperture contraction rate in N. bachei is approximately 20% of
the average jet velocity. Broken lines indicate measurement uncertainty of
±10% associated with calculation of vortex formation time (see
Materials and methods). Note that the local plateau in the vortex formation
time near t/Tejection=0.7 is within the error of
the measurement trend.
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Fig. 4. Optimal vortex formation during fast-swimming of N. bachei. Image
from dye flow visualization showing vortex formation in the wake of the
animal. Image height is 7.2 mm. No trailing flow exists directly behind the
vortex (see also movie in supplementary material), supporting the conclusion
that the animal generates a single vortex (shown schematically in inset) per
swimming cycle, despite the fact that
T*max 8.
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Fig. 5. Measurement and models of N. bachei swimming trajectory. The ratio
of instantaneous forward motion (relative to its position at the start of the
measured swimming cycle) to the maximum forward motion,
x/Xmax, is plotted versus the current point in
the full swimming cycle (i.e. ejection and relaxation). Both the real animal
(solid black line) and the model that includes transient velar kinematics
(broken black line) achieve continuous forward motion, whereas the model that
assumes a constant velar aperture (dotted grey line) swims backward during the
latter portion of the swimming cycle. Measurement uncertainty is
±3%.
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© The Company of Biologists Ltd 2006