Cyprinid swimming behaviour in response to turbulent flow

a b s t r a c t Turbulence is a complex phenomenon which commonly occurs in river and fishway flows. It is a diffi- cult subject to study, especially biologically, yet turbulence may affect fish movements and fish passage efficiency. Studies on quantifying fish responses to turbulence, particularly within fishways, are lacking. This study investigated the swimming behaviour of 140 adult Iberian barbel (Luciobarbus bocagei) of two size-classes (small fish: 15 ≤ TL < 25 cm, large fish: 25 < TL ≤ 35 cm) under turbulent flow conditions cre- ated by three submerged orifice arrangements in an experimental pool-type fishway: (i) offset orifices, (ii) straight orifices and (iii) straight orifices with a deflector bar of 0.5bo located at 0.2L from the inlet orifices, where bo is the width of the square orifices ranging from 0.18 to 0.23 m and L is the pool length (1.90 m). Water velocity and turbulence (turbulent kinetic energy, Reynolds shear stress, turbulence intensity and eddy size) were characterized using a 3D Acoustic Doppler Velocimeter (ADV) and were related with fish swimming behaviour. The influence of turbulent flow on the swimming behaviour of barbel was assessed through the number of successful fish passage attempts and associated passage times. The amount of time fish spent in a certain cell of the pool (transit time) was measured and related to hydraulic conditions. The highest rates of passage and the corresponding lowest times were found in experiments conducted with offset orifices. Although size-related behavioural responses to turbulence were observed, Reynolds shear stress appeared as one of the most important turbulence descriptors explaining fish transit time for both size-classes in experiments conducted with offset and straight orifices; furthermore, swimming behaviour of larger fish was found to be strongly affected by the eddies created, in particular by those of similar size to fish total length, which were mainly found in straight orifices with a deflector bar arrange- ment. The results provide valuable insights on barbel swimming behavioural responses to turbulence, which may help engineers and biologists to develop effective systems for the passage of this species and others with similar biomechanical capacities.

[1]  A.M.O. Smith,et al.  Turbulence models and their application in hydraulics: W. Rodi, University of Karlsruhe, International Association for Hydraulic Research, Rotterdamseweg 185, 2600 MH Delft, The Netherlands , 1981 .

[2]  António N. Pinheiro,et al.  Passage efficiency of offset and straight orifices for upstream movements of Iberian barbel in a pool‐type fishway , 2012 .

[3]  David L. Smith,et al.  Response of juvenile rainbow trout to turbulence produced by prismatoidal shapes , 2005 .

[4]  Paul W. Webb,et al.  Simple Physical Principles and Vertebrate Aquatic Locomotion , 1988 .

[5]  M. Lucas,et al.  Effects of a flow‐gauging weir on the migratory behaviour of adult barbel, a riverine cyprinid , 1997 .

[6]  J. Durbin,et al.  Testing for serial correlation in least squares regression. II. , 1950, Biometrika.

[7]  Paul W. Webb,et al.  Do Brown Trout Choose Locations with Reduced Turbulence , 2006 .

[8]  A. Roy,et al.  A model of total swimming costs in turbulent flow for juvenile Atlantic salmon (Salmo salar) , 2005 .

[9]  Bahman Shafii,et al.  Use of the Average and Fluctuating Velocity Components for Estimation of Volitional Rainbow Trout Density , 2006 .

[10]  Milo C Bell,et al.  Fisheries Handbook of Engineering Requirements and Biological Criteria , 1990 .

[11]  M. Magalhães,et al.  Riparian ecotones and spatial variation of fish assemblages in Portuguese lowland streams , 1995, Hydrobiologia.

[12]  Christos Katopodis,et al.  Developing a toolkit for fish passage, ecological flow management and fish habitat works , 2005 .

[13]  Martin Wikelski,et al.  Migration by soaring or flapping: numerical atmospheric simulations reveal that turbulence kinetic energy dictates bee-eater flight mode , 2011, Proceedings of the Royal Society B: Biological Sciences.

[14]  J. Liao,et al.  A review of fish swimming mechanics and behaviour in altered flows , 2007, Philosophical Transactions of the Royal Society B: Biological Sciences.

[15]  Charles H. Clay,et al.  Design of Fishways and Other Fish Facilities , 1995 .

[16]  Fao Dvwk.  Fish Passes: Design, Dimensions, and Monitoring , 2002 .

[17]  Ian G. Cowx,et al.  Rehabilitation of rivers for fish , 1998 .

[18]  D. S. Pavlov,et al.  The Effects of Flow Turbulence on the Behavior and Distribution of Fish , 2000 .

[19]  J. Armstrong,et al.  Movements of Atlantic salmon migrating upstream through a fish‐pass complex in Scotland , 2003 .

[20]  F. Capel,et al.  Desarrollo de curvas de preferencia de microhábitat para Leuciscus pyrenaicus y Barbus bocagei por buceo en el río Jarama (Cuenca del Tajo). , 1999 .

[21]  N. Rajaratnam,et al.  Hydraulics of simple habitat structures , 2001 .

[22]  P W Webb,et al.  Turbulence: does vorticity affect the structure and shape of body and fin propulsors? , 2010, Integrative and comparative biology.

[23]  G. Lauder,et al.  Fish Exploiting Vortices Decrease Muscle Activity , 2003, Science.

[24]  John Guckenheimer,et al.  Paddling mode of forward flight in insects. , 2011, Physical review letters.

[25]  A. Roy,et al.  The effect of turbulence on the cost of swimming for juvenile Atlantic salmon (Salmo salar) , 2003 .

[26]  Christoph Brücker,et al.  Entraining in trout: a behavioural and hydrodynamic analysis , 2010, Journal of Experimental Biology.

[27]  Mufeed. Odeh,et al.  Evaluation of the Effects of Turbulence on the Behavior of Migratory Fish, 2002 Final Report. , 2002 .

[28]  B. Quintella,et al.  The critical swimming speed of Iberian barbel Barbus bocagei in relation to size and sex , 2008 .

[29]  M. T. Ferreira,et al.  Effects of water velocity and turbulence on the behaviour of Iberian barbel (Luciobarbus bocagei, Steindachner 1864) in an experimental pool‐type fishway , 2011 .

[30]  A. I. Lupandin.  Effect of Flow Turbulence on Swimming Speed of Fish , 2005, Biology Bulletin.

[31]  P. Bradshaw,et al.  Turbulence Models and Their Application in Hydraulics. By W. RODI. International Association for Hydraulic Research, Delft, 1980. Paperback US $15. , 1983, Journal of Fluid Mechanics.

[32]  E. Baras,et al.  A comprehensive assessment of the failure of Barbus barbus spawning migrations through a fish pass in the canalized River Meuse (Belgium) , 1994 .

[33]  M. T. Ferreira,et al.  Efficacy of a nature‐like bypass channel in a Portuguese lowland river , 2005 .

[34]  R. J. Smith.  Social behaviour, homing and migration , 1991 .

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

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

[37]  James W. A. Grant,et al.  A quantitative assessment of fish passage efficiency , 2012 .

[38]  M. A. Skorobogatov,et al.  The critical current velocity of fish and the degree of flow turbulence , 1982 .

[39]  F S Hover,et al.  Vorticity Control in Fish-like Propulsion and Maneuvering1 , 2002, Integrative and comparative biology.

[40]  H. Tritico,et al.  The effects of turbulent eddies on the stability and critical swimming speed of creek chub (Semotilus atromaculatus) , 2010, Journal of Experimental Biology.

[41]  Paul W. Webb,et al.  Response latencies to postural disturbances in three species of teleostean fishes , 2004, Journal of Experimental Biology.

[42]  M. Eens,et al.  Performance of a pool-and-weir fish pass for small bottom-dwelling freshwater fish species in a regulated lowland river , 2007 .

[43]  J. Meeuwig,et al.  Freshwater Protected Areas: Strategies for Conservation , 2002, Conservation biology : the journal of the Society for Conservation Biology.

[44]  M. Tachie,et al.  Proper Orthogonal Decomposition Analysis of Separated and Reattached Pressure Gradient Turbulent Flows , 2009 .

[45]  Francisco Martínez-Capel,et al.  Microhabitat use by three endemic Iberian cyprinids in Mediterranean rivers (Tagus River Basin, Spain) , 2009 .

[46]  Marshall C. Richmond,et al.  Exploring the Role of Shear Stress and Severe Turbulence in Downstream Fish Passage , 1999 .

[47]  Anders Hedenström,et al.  Migration by soaring or flapping flight in birds: the relative importance of energy cost and speed , 1993 .

[48]  J. Lumley,et al.  A First Course in Turbulence , 1972 .

[49]  P. Magnan,et al.  Relationship between individual variation in morphological characters and swimming costs in brook charr (Salvelinus fontinalis) and yellow perch (Perca flavescens). , 2002, The Journal of experimental biology.

[50]  P. Webb,et al.  Control of Posture, Depth, and Swimming Trajectories of Fishes1 , 2002, Integrative and comparative biology.

[51]  N. Rajaratnam,et al.  Flow regimes and structure in pool and weir fishways , 2004 .