Volumetric imaging of shark tail hydrodynamics reveals a three-dimensional dual-ring vortex wake structure

Understanding how moving organisms generate locomotor forces is fundamental to the analysis of aerodynamic and hydrodynamic flow patterns that are generated during body and appendage oscillation. In the past, this has been accomplished using two-dimensional planar techniques that require reconstruction of three-dimensional flow patterns. We have applied a new, fully three-dimensional, volumetric imaging technique that allows instantaneous capture of wake flow patterns, to a classic problem in functional vertebrate biology: the function of the asymmetrical (heterocercal) tail of swimming sharks to capture the vorticity field within the volume swept by the tail. These data were used to test a previous three-dimensional reconstruction of the shark vortex wake estimated from two-dimensional flow analyses, and show that the volumetric approach reveals a different vortex wake not previously reconstructed from two-dimensional slices. The hydrodynamic wake consists of one set of dual-linked vortex rings produced per half tail beat. In addition, we use a simple passive shark-tail model under robotic control to show that the three-dimensional wake flows of the robotic tail differ from the active tail motion of a live shark, suggesting that active control of kinematics and tail stiffness plays a substantial role in the production of wake vortical patterns.

[1]  A. Hedenström,et al.  Bat Flight Generates Complex Aerodynamic Tracks , 2007, Science.

[2]  E. Longmire,et al.  Vortex dynamics in jets from inclined nozzles , 1997 .

[3]  C. A. Pell,et al.  Mechanical control of swimming speed: stiffness and axial wave form in undulating fish models , 1995, The Journal of experimental biology.

[4]  George V. Lauder,et al.  Low-dimensional models and performance scaling of a highly deformable fish pectoral fin , 2009, Journal of Fluid Mechanics.

[5]  Performance prediction of point-based three-dimensional volumetric measurement systems , 2008 .

[6]  George V Lauder,et al.  The mechanics of active fin-shape control in ray-finned fishes , 2007, Journal of The Royal Society Interface.

[7]  G. V. Lauder,et al.  Biomechanics: Hydrodynamic function of the shark's tail , 2004, Nature.

[8]  R. Alexander,et al.  THE LIFT PRODUCED BY THE HETEROCERCAL TAILS OF SELACHII , 1965 .

[9]  George V. Lauder,et al.  Function of the Caudal Fin During Locomotion in Fishes: Kinematics, Flow Visualization, and Evolutionary Patterns1 , 2000 .

[10]  Eric D. Tytell,et al.  Median fin function in bluegill sunfish Lepomis macrochirus: streamwise vortex structure during steady swimming , 2006, Journal of Experimental Biology.

[11]  George V. Lauder,et al.  Hydrodynamics of Undulatory Propulsion , 2005 .

[12]  George V Lauder,et al.  Volumetric imaging of fish locomotion , 2011, Biology Letters.

[13]  G. Lauder,et al.  The hydrodynamics of eel swimming , 2004, Journal of Experimental Biology.

[14]  A Hedenström,et al.  Vortex wakes generated by robins Erithacus rubecula during free flight in a wind tunnel , 2006, Journal of The Royal Society Interface.

[15]  Haibo Dong,et al.  Locomotion with flexible propulsors: I. Experimental analysis of pectoral fin swimming in sunfish , 2006, Bioinspiration & biomimetics.

[16]  M. Triantafyllou,et al.  An Efficient Swimming Machine , 1995 .

[17]  J.-M. Miao,et al.  Effect of flexure on aerodynamic propulsive efficiency of flapping flexible airfoil , 2006 .

[18]  Morteza Gharib,et al.  Flow patterns generated by oblate medusan jellyfish: field measurements and laboratory analyses , 2005, Journal of Experimental Biology.

[19]  B. Tobalske,et al.  Aerodynamics of the hovering hummingbird , 2005, Nature.

[20]  Bret W Tobalske,et al.  Aerodynamics of wing-assisted incline running in birds , 2007, Journal of Experimental Biology.

[21]  A. Smits,et al.  Thrust production and wake structure of a batoid-inspired oscillating fin , 2005, Journal of Fluid Mechanics.

[22]  George V Lauder,et al.  Experimental Hydrodynamics and Evolution: Function of Median Fins in Ray-finned Fishes1 , 2002, Integrative and comparative biology.

[23]  J. Dabiri,et al.  Fast-swimming hydromedusae exploit velar kinematics to form an optimal vortex wake , 2006, Journal of Experimental Biology.

[24]  G.V. Lauder,et al.  Morphology and experimental hydrodynamics of fish fin control surfaces , 2004, IEEE Journal of Oceanic Engineering.

[25]  M. Triantafyllou,et al.  Wake mechanics for thrust generation in oscillating foils , 1991 .

[26]  Brooke E Flammang,et al.  Functional morphology of the radialis muscle in shark tails , 2009, Journal of morphology.

[27]  Ellen K. Longmire,et al.  Volumetric velocity measurements of vortex rings from inclined exits , 2010 .

[28]  York Winter,et al.  The near and far wake of Pallas' long tongued bat (Glossophaga soricina) , 2008, Journal of Experimental Biology.

[29]  G. Lauder,et al.  Function of the heterocercal tail in white sturgeon: flow visualization during steady swimming and vertical maneuvering. , 2000, The Journal of experimental biology.

[30]  S. R. Olsen,et al.  An in situ rapid heat–quench cell for small-angle neutron scattering , 2008 .

[31]  Lauder,et al.  Heterocercal tail function in leopard sharks: a three-dimensional kinematic analysis of two models , 1996, The Journal of experimental biology.

[32]  George V Lauder,et al.  Hydrodynamics of caudal fin locomotion by chub mackerel, Scomber japonicus (Scombridae). , 2002, The Journal of experimental biology.

[33]  George V Lauder,et al.  Advances in comparative physiology from high-speed imaging of animal and fluid motion. , 2008, Annual review of physiology.

[34]  A Hedenström,et al.  A family of vortex wakes generated by a thrush nightingale in free flight in a wind tunnel over its entire natural range of flight speeds , 2003, Journal of Experimental Biology.

[35]  J. Videler,et al.  Fish foot prints: morphology and energetics of the wake behind a continuously swimming mullet (Chelon labrosus Risso). , 1997, The Journal of experimental biology.

[36]  M. Dickinson,et al.  Spanwise flow and the attachment of the leading-edge vortex on insect wings , 2001, Nature.

[37]  G. Lauder,et al.  Hydrodynamic function of dorsal and anal fins in brook trout (Salvelinus fontinalis) , 2007, Journal of Experimental Biology.

[38]  J. R. Simons The Direction of the Thrust Produced by the Heterocercal Tails of Two Dissimilar Elasmobranchs: The Port Jackson Shark, Heterodontus Portusjacksoni (Meyer), and the Piked Dogfish, Squalus Megalops (Macleay) , 1970 .

[39]  Keith Stewart Thomson,et al.  Body Form and Locomotion in Sharks , 1977 .

[40]  Rajat Mittal,et al.  The effect of fin ray flexural rigidity on the propulsive forces generated by a biorobotic fish pectoral fin , 2010, Journal of Experimental Biology.

[41]  Joseph Katz,et al.  Hydrodynamic propulsion by large amplitude oscillation of an airfoil with chordwise flexibility , 1978, Journal of Fluid Mechanics.

[42]  James Tangorra,et al.  Fish biorobotics: kinematics and hydrodynamics of self-propulsion , 2007, Journal of Experimental Biology.

[43]  G. Lauder,et al.  Function of the heterocercal tail in sharks: quantitative wake dynamics during steady horizontal swimming and vertical maneuvering. , 2002, The Journal of experimental biology.

[44]  Francisco Pereira,et al.  Defocusing digital particle image velocimetry: a 3-component 3-dimensional DPIV measurement technique. Application to bubbly flows , 2000 .

[45]  E. Longmire,et al.  Vortex rings from cylinders with inclined exits , 1998 .

[46]  A. Maia,et al.  Biomechanics of Locomotion in Sharks, Rays, and Chimeras , 2012 .