The functional role of caudal and anal/dorsal fins during the C-start of a bluegill sunfish

SUMMARY Fast starts are crucial in the survival of aquatic swimmers to capture prey or avoid predators. Currently, it is widely accepted that during C-starts: (1) the caudal fin generates a considerable hydrodynamic force; and (2) anal/dorsal fins are erected to significantly increase the hydrodynamic force. In this work, the above hypotheses on the role of fins during C-starts are studied using experimentally guided numerical simulations of four bluegill sunfish, whose fins are removed or erected. The amount of force created by the body and fins at each time instant was not constant and varied during the C-start. Nevertheless, in agreement with hypothesis (1), up to 70% of the instantaneous hydrodynamic force was generated by the tail during Stage 2 of the C-start, when the sunfish rapidly bends out of the C-shape. Additionally, the contribution in Stage 1, when the sunfish bends into a C-shape, is less than 20% at each instant. Most of the force in Stage 1 was produced by the body of the sunfish. In contrast to hypothesis (2), the effect of erection/removal of the fins was less than 5% of the instantaneous force in both Stages 1 and 2, except for a short period of time (2 ms) just before Stage 2. However, it is known that the anal/dorsal fins are actively controlled during the C-start from muscle activity measurements. Based on the results presented here, it is suggested that the active control of the anal/dorsal fins can be related to retaining the stability of the sunfish against roll and pitch movements during the C-start. Furthermore, the erection of the fins increases the moment of inertia to make the roll and pitch movements more difficult.

[1]  Triantafyllou,et al.  Near-body flow dynamics in swimming fish , 1999, The Journal of experimental biology.

[2]  David N. Reznick,et al.  Do faster starts increase the probability of evading predators , 2005 .

[3]  Robert W. Blake,et al.  Energetics of piscivorous predator-prey interactions , 1988 .

[4]  P. Webb,et al.  Strike tactics of Esox. , 1980, Canadian journal of zoology.

[5]  Matthew Ringuette,et al.  Vortex formation and saturation for low-aspect-ratio rotating flat-plate fins , 2012 .

[6]  Robert W. Blake,et al.  ESCAPE TRAJECTORIES IN ANGELFISH (PTEROPHYLLUM EIMEKEI) , 1993 .

[7]  G. Lauder,et al.  Hydrodynamics of the escape response in bluegill sunfish, Lepomis macrochirus , 2008, Journal of Experimental Biology.

[8]  A. Kinkhabwala,et al.  Mapping a sensory-motor network onto a structural and functional ground plan in the hindbrain , 2011, Proceedings of the National Academy of Sciences.

[9]  G. Lauder,et al.  Function of the dorsal fin in bluegill sunfish: Motor patterns during four distinct locomotor behaviors , 1996, Journal of morphology.

[10]  Robert W. Blake,et al.  On the error involved in high-speed film when used to evaluate maximum accelerations of fish , 1989 .

[11]  Spierts,et al.  Kinematics and muscle dynamics of C- and S-starts of carp (Cyprinus carpio L.). , 1999, The Journal of experimental biology.

[12]  Hale,et al.  Mechanics of the fast-start: muscle function and the role of intramuscular pressure in the escape behavior of amia calva and polypterus palmas , 1998, The Journal of experimental biology.

[13]  E. Standen,et al.  Escape manoeuvres in the spiny dogfish (Squalus acanthias) , 2004, Journal of Experimental Biology.

[14]  John O. Dabiri,et al.  On the estimation of swimming and flying forces from wake measurements , 2005, Journal of Experimental Biology.

[15]  Robert W. Blake,et al.  The Kinematics and Performance of the Escape Response in the Angelfish (Pterophyllum Eimekei) , 1991 .

[16]  G. Dehnhardt,et al.  Hydrodynamic determination of the moving direction of an artificial fin by a harbour seal (Phoca vitulina) , 2010, Journal of Experimental Biology.

[17]  Wolf Hanke,et al.  The hydrodynamic trails of Lepomis gibbosus (Centrarchidae), Colomesus psittacus (Tetraodontidae) and Thysochromis ansorgii (Cichlidae) investigated with scanning particle image velocimetry , 2004, Journal of Experimental Biology.

[18]  Brad A. Chadwell,et al.  Musculoskeletal morphology and regionalization within the dorsal and anal fins of bluegill sunfish (Lepomis macrochirus) , 2012, Journal of morphology.

[19]  George V Lauder,et al.  Median fin function during the escape response of bluegill sunfish (Lepomis macrochirus). II: Fin-ray curvature , 2012, Journal of Experimental Biology.

[20]  P. Webb Body and Fin Form and Strike Tactics of Four Teleost Predators Attacking Fathead Minnow (Pimephales promelas) Prey , 1984 .

[21]  F. Rank,et al.  Balancing Requirements for Stability and Maneuverability in Cetaceans ] , 2002 .

[22]  J. Fetcho Spinal Network of the Mauthner Cell (Part 1 of 2) , 1991 .

[23]  Fotis Sotiropoulos,et al.  A numerical method for solving the 3D unsteady incompressible Navier-Stokes equations in curvilinear domains with complex immersed boundaries , 2007, J. Comput. Phys..

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

[25]  C A Bergstrom,et al.  Fast-start swimming performance and reduction in lateral plate number in threespine stickleback , 2002 .

[26]  D. Weihs,et al.  A hydrodynamical analysis of fish turning manoeuvres , 1972, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[27]  Johnston,et al.  The biomechanics of fast-starts during ontogeny in the common carp cyprinus carpio , 1999, The Journal of experimental biology.

[28]  Ulrike K Müller,et al.  Flow patterns of larval fish: undulatory swimming in the intermediate flow regime , 2008, Journal of Experimental Biology.

[29]  R. W. Blake,et al.  Mechanics of the startle response in the northern pike, Esox lucius , 1991 .

[30]  M S Triantafyllou,et al.  A fast-starting mechanical fish that accelerates at 40 m s−2 , 2010, Bioinspiration & biomimetics.

[31]  J M Wakeling,et al.  Muscle power output limits fast-start performance in fish. , 1998, The Journal of experimental biology.

[32]  P. Moin,et al.  Eddies, streams, and convergence zones in turbulent flows , 1988 .

[33]  Paul W. Webb,et al.  Fast-start Performance and Body Form in Seven Species of Teleost Fish , 1978 .

[34]  D. Ellerby,et al.  Spatial variation in fast muscle function of the rainbow trout Oncorhynchus mykiss during fast-starts and sprinting. , 2001, The Journal of experimental biology.

[35]  Yousef Saad,et al.  Iterative methods for sparse linear systems , 2003 .

[36]  Paul W. Webb,et al.  EFFECTS OF MEDIAN-FIN AMPUTATION ON FAST-START PERFORMANCE OF RAINBOW TROUT (SALMO GAIRDNERI) , 1977 .

[37]  G. Lauder,et al.  Dorsal and anal fin function in bluegill sunfish Lepomis macrochirus: three-dimensional kinematics during propulsion and maneuvering , 2005, Journal of Experimental Biology.

[38]  Paul W. Webb,et al.  Does body and fin form affect the maneuverability of fish traversing vertical and horizontal slits? , 2004, Environmental Biology of Fishes.

[39]  I. Borazjani,et al.  Numerical investigation of the hydrodynamics of anguilliform swimming in the transitional and inertial flow regimes , 2009, Journal of Experimental Biology.

[40]  Fotis Sotiropoulos,et al.  Curvilinear immersed boundary method for simulating fluid structure interaction with complex 3D rigid bodies , 2008, J. Comput. Phys..

[41]  H. Bleckmann,et al.  Hydrodynamic Trail-Following in Harbor Seals (Phoca vitulina) , 2001, Science.

[42]  George V Lauder,et al.  Median fin function during the escape response of bluegill sunfish (Lepomis macrochirus). I: Fin-ray orientation and movement , 2012, Journal of Experimental Biology.

[43]  Domenici,et al.  The kinematics and performance of fish fast-start swimming , 1997, The Journal of experimental biology.

[44]  R. C. Eaton,et al.  The Mauthner cell and other identified neurons of the brainstem escape network of fish , 2001, Progress in Neurobiology.

[45]  I. Borazjani,et al.  Numerical investigation of the hydrodynamics of carangiform swimming in the transitional and inertial flow regimes , 2008, Journal of Experimental Biology.

[46]  Brenden P. Epps,et al.  Impulse generated during unsteady maneuvering of swimming fish , 2007 .

[47]  T. Lingham‐Soliar Dorsal fin in the white shark, Carcharodon carcharias: A dynamic stabilizer for fast swimming , 2005, Journal of morphology.

[48]  G. V. Lauder,et al.  Red and white muscle activity and kinematics of the escape response of the bluegill sunfish during swimming , 1993, Journal of Comparative Physiology A.

[49]  R. Blake Fish functional design and swimming performance , 2004 .

[50]  J. Fetcho,et al.  Spinal network of the Mauthner cell. , 1991, Brain, behavior and evolution.

[51]  M. Lighthill Large-amplitude elongated-body theory of fish locomotion , 1971, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[52]  Fotis Sotiropoulos,et al.  Hydrodynamics of the bluegill sunfish C-start escape response: three-dimensional simulations and comparison with experimental data , 2012, Journal of Experimental Biology.

[53]  J. M. Wakeling,et al.  Predicting muscle force generation during fast-starts for the common carp Cyprinus carpio , 1999, Journal of Comparative Physiology B.

[54]  F. Sotiropoulos,et al.  A hybrid Cartesian/immersed boundary method for simulating flows with 3D, geometrically complex, moving bodies , 2005 .

[55]  I. Borazjani,et al.  On the role of form and kinematics on the hydrodynamics of self-propelled body/caudal fin swimming , 2010, Journal of Experimental Biology.

[56]  B. Prendergast,et al.  Comparative studies on the Mauthner cell of teleost fish in relation to sensory input. , 1995, Brain, behavior and evolution.

[57]  George V Lauder,et al.  The C-start escape response of Polypterus senegalus: bilateral muscle activity and variation during stage 1 and 2. , 2002, The Journal of experimental biology.

[58]  Jeffrey A. Walker,et al.  ESTIMATING VELOCITIES AND ACCELERATIONS OF ANIMAL LOCOMOTION: A SIMULATION EXPERIMENT COMPARING NUMERICAL DIFFERENTIATION ALGORITHMS , 1998 .

[59]  J. Katz,et al.  On the role of copepod antennae in the production of hydrodynamic force during hopping , 2010, Journal of Experimental Biology.

[60]  Paolo Domenici,et al.  Escape behaviour of solitary herring (Clupea harengus ) and comparisons with schooling individuals , 1997 .

[61]  Paul W. Webb,et al.  Stability and Maneuverability , 2005 .

[62]  G. J. Rose,et al.  Activation of Mauthner neurons during prey capture , 1993, Journal of Comparative Physiology A.

[63]  Eric D. Tytell,et al.  Do trout swim better than eels? Challenges for estimating performance based on the wake of self-propelled bodies , 2007 .

[64]  D. Weihs,et al.  The mechanism of rapid starting of slender fish. , 1973, Biorheology.

[65]  Robert W. Blake,et al.  Prey capture and the fast-start performance of northern pike Esox lucius , 1991 .

[66]  Petros Koumoutsakos,et al.  C-start: optimal start of larval fish , 2012, Journal of Fluid Mechanics.

[67]  Daniel Weihs,et al.  Stability Versus Maneuverability in Aquatic Locomotion1 , 2002, Integrative and comparative biology.