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Locomotion in the Biology of Large Aquatic Vertebrates

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Locomotion in the Biology of Large Aquatic Vertebrates Transactions of the American Fisheries Society 119:629-641. 1990 <S> Copyright by the American Fisheries Society 1990 Locomotio Biologe th n i f Largyo e Aquatic Vertebrates PAU . WEBLW B Department of Biology Schooland of Natural Resources University of Michigan. Ann Arbor, Michigan 48109-1115. USA. and Ministry of Agriculture. Fisheries Food.and Fisheries Laboratory Lowestoft. Suffolk NR33 OHT, England VIVIAN DE BUFFR£NIL Museum National d'Histoire Naturelle. 75005 Paris. France Abstract.—As aquatic vertebrates increase in size, hydrofoils, which use lift to generate thrust, increasingle ar y use propulsorss da factoe On . r affectin magnitude gth life t th are e forcf eo th a s ei of the propulsor. Resistance to cruising and sprints is mainly due to drag, but inertia is important during maneuvers when animals accelerate or turn. The inertia of the body and entrained water, which is proportional to body volume, resists acceleration. Because a thrust that is proportional to surface are s usei a maneuveo dt a resistancr e tha s proportionai t volumeo t l , acceleration performance and maneuverability are expected to decline with increasing size. This trend is ame- liorated to some extent by the high swimming speeds attainable by warm-bodied vertebrates and the reduced resistanc acceleratioo et n characteristi skeletone th f co dolphinf so ichthyosaursd san . Maneuvers are essential for capture of elusive prey and avoidance of predators. As they increase in size, aquatic vertebrates use various means to ensure that their prey are less maneuverable than they. These include consumption of increasingly smaller prey relative to predator body size (cul- minatin filten gi r feedin largese th y gb t aquatic vertebrates); behavior concentrateo st , disturbd an , disorient prey; and ambushing or suction feeding that avoid whole-body acceleration. Advantages of warm muscles are seen in the ability of endotherms to take more maneuverable prey than can ectotherms of the same size. Young stages of large aquatic vertebrates could be especially vulnerable predatorso t ; viviparit r spawninyo productivn gi e patches provide r rapifo s d growth through vulnerable stages. All organisms must function within limits im- that equal totae th s l resistanc bode d th yan f o e posed by laws that describe the properties and appendages at every instant. Both thrust and re- interaction f matteso r (Danie Webd an l b 1987). sistance arise from transfer f momentuso m be- These restric range th t optionf eo forr d fo smtwee an watere animae th nth d . an Thesl e transfers function as animal performance approaches the are described in terms of a small set of mecha- boundarie physicay b t sse l laws. Thus, physical nisms: friction drag, pressure drag, accelliftd an , - principle usee b predico dt n stesca d an tt ideas, eration reaction (Danie Webd an l b 1987; Webb leadin fulleo gt r understandin biologe th f g o f yo 1988). organisms (Bartholomew 1987; Danie Webd an l b Frictio r viscouo n s drag arises from velocity 1987; Webb 1988) apple W . y this approac o ht gradient whicn i s h viscosity resist e relativth s e consider the consequences of large size for loco- motion of adjacent water molecules. For verte- motor capabilities of aquatic vertebrates. We first brates, large velocity gradients and viscous forces discuss mechanical concepts and forces acting on typically occur in a thin layer—the boundary lay- swimmers. These set constraints on function, of er—on body and propulsor surfaces. The viscous whic considee hw r maneuverabilit e mosb o ytt forc s proportionai e surfaco t e l th o et ared an a important. We then explore the implications of square of speed. these constraints for aquatic vertebrates in terms Pressure drag occurs because the flow pattern of muscle temperature regulation, morpholog d ydifferan s upstrea downstread man solia mf o d object anatomy, predator-prey interactions, foraging sucvertebrata s ha e bodflaa r ty o surfac e that patterns lifd ean , history. move r wits inciden e oa lik hth n ea t flow normal surfacee th boundaro e t .Th y layer eventually sep- . _, arates from the surface of most vertebrate-sized m r oorceo s object created san waksa e downstream prese Th . - Aquatic vertebrates use the tail and other ap- sure in the wake is lower than the pressure up- pendages to swim. These propulsors generate thrust stream, and the resulting pressure difference re- 629 630 WEB BUFFRENID BAN L tards forward motion. Streamlining delays <»}L/u= a (1) separation of the boundary layer from body until and downstrea close is ti th eo t m (trailing o p bodyfa ) ti , hencd an e minimizes pressure drag. Conversely, Re Lulv,= (2) ensurer anoa s that separation occurs early, which L = characteristic length, usually total length; maximizes pressure drag. In either situation, pres- velocity= u ; sure drag is proportional to surface area and the v = kinematic viscosity of water; squar speedf eo . a= radia) n frequency, rf\ Liftforce th , e generate winga y db , also arises oscillatio= f n frequency. from a pressure difference between opposite sur- faces of an object. In this case, the object moves Reynolde Th s numbe largr rfo e vertebrates typ- at a small angle to the incident flow, and the ically exceeds 10s. As a result, drag, lift, and ac- boundary layer separates fro e surfacmth e only celeration reactio dominane th e nar t forces acting close to the trailing edge. Water is accelerated over bode opropulsorsd nth yan relative Th . e impor- one surface, resulting in a lower pressure than on steade tanc th unsteadd f eyo an y force compo- the other surface. The pressure difference Gift) is nent estimates si Whe. a y objecdn b na t oscillates oriented approximately normal to the incident many times during the time a particle of water flow, and its magnitude is proportional to surface traverses the object's length, or chord, that water area and the square of speed. particle is accelerated and decelerated during its Acceleration of an object dissipates energy by passage. The largs i acceleratio d na ean n reaction inducing acceleration of the water in the vicinity force e largear s . Hydromechanical analyses of the body. This energy can be visualized as a (Lighthill 1975; Daniel 1984) show this occurs resistance increment, the acceleration reaction, when a exceeds 0.4. Conversely, if an object os- equated to that of an added mass of water accel- cillates slowly thao s , particlta watef eo r travers- erated with the object. Acceleration resistance is t inacceleratechore gno th s di d much fros mit hence proportional to acceleration rate and mass. steade path th smals i d ya , an lforce s dominate. Several classification f theso s e n i force e ar s This occurs whe less i nsa than 0.1. Both steady common use, linking force termn i s f shareso d unsteadd an y force components shoul takee db n variables that affect their respective magnitudes. int (Danie4 o 0. accoun Webd < lan a 0.r b1< tfo Acceleration reactio proportionans i masso lt d ,an 1987). hence volume t alsi calles o i s , dvolumea force. For propulsors, a decreases with increasing an- It is also proportional to acceleration, or the rate imal size. Thus/for oscillating propulsor pros si - of change of velocity with time. Therefore, accel- portional to L-°45 to L-1 (Peters 1983; Caider eration reaction is defined as an unsteady force. 1984; Heglund and Taylor 1988). As a result, <r Dra Hd proportionae Agan ar surface th o t l e area approaches 0.1 for propulsors of large vertebrates of an object and are called surface forces. In ad- (Daniel and Webb 1987). Steady forces also be- dition, the functione yar f speedo s theo s , y also come larger relative to unsteady forces as ampli- callee ar d steady forces. tude increases. The amplitudes of propulsor mo- Animals may employ drag, lift, and acceleration tions increase roughly wit (PeterhL s 1983), which reactio generato nt e thrust. Eac f thesho e forces further increase importance sth steadf eo y forces may also contribute to resistance, although lift for propulsor largef so r organisms. probabl uses yresistanca i s da e force only during The body does not oscillate as much as the tail, braking (see Danie Webd lan b 1987; Webb 1988). and tuna-shaped vertebrate visualizee b n sca s da magnitude Th relativd ean e importanc thesf eo e a propulsor attached to a more-or-less rigid body forces vary with flow conditions. Whe animan na l (Lighthill 1977). The bode th nr verys i < fo r y much swims well belo water/aie wth r surface, flow con- smaller than 0.1. ditions are related to reduced frequency (a) and Thus draexpectee lifd ar tgan increase b o dt - Reynolds number (Re). Reduced frequency indi- ingly important for thrust and resistance as aquat- cate extene sth whico t h oscillation objectn a f so , ic vertebrates increase in size because Re is high such as a propulsor, affect the flow of a fluid pass- and a becomes small. Acceleration reaction will ing over that object. Reynolds number indicates still have an effect, mainly for the propulsors of the relative importance of pressure drag, lift, and medium-sized vertebrates (Daniel and Webb acceleration reaction compared to viscous forces. 1987), but energy expended in overcoming these These terms are formally defined as forces more commonly contribute o energt s y LOCOMOTIO LARGF NO E AQUATIC VERTEBRATES 631 efficiencwastagw lo d ean y (Danie Webd lan b 1987; Large aspect ratio and tapering of a hydrofoil to- Webb 1988).
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