Experimental hydrodynamics of fish locomotion: functional insights from wake visualization

Despite enormous progress during the last twenty years in understandingthe mechanistic basis of aquatic animal propulsion—a taskinvolving the construction of a substantial data base on patternsof fin and body kinematics and locomotor muscle function—thereremains a key area in which biologists have little information:the relationship between propulsor activity and water movementin the wake. How is internal muscular force translated intoexternal force exerted on the water? What is the pattern offluid force production by different fish fins (e.g., pectoral,caudal, dorsal) and how does swimming force vary with speedand among species? These types of questions have received considerableattention in analyses of terrestrial locomotion where forceoutput by limbs can be measured directly with force plates.But how can forces exerted by animals moving through fluid bemeasured? The advent of digital particle image velocimetry (DPIV)has provided an experimental hydrodynamic approach for quantifyingthe locomotor forces of freely moving animals in fluids, andhas resulted in significant new insights into the mechanismsof fish propulsion. In this paper we present ten "lessons learned"from the application of DPIV to problems of fish locomotionover the last five years. (1) Three-dimensional DPIV analysisis critical for reconstructing wake geometry. (2) DPIV analysisreveals the orientation of locomotor reaction forces. (3) DPIVanalysis allows calculation of the magnitude of locomotor forces.(4) Swimming speed can have a major impact on wake structure.(5) DPIV can reveal interspecific differences in vortex wakemorphology. (6) DPIV analysis can provide new insights intothe limits to locomotor performance. (7) DPIV demonstrates thefunctional versatility of fish fins. (8) DPIV reveals hydrodynamicforce partitioning among fins. (9) DPIV shows that wake interactionamong fins may enhance thrust production. (10) Experimentalhydrodynamic analysis can provide insight into the functionalsignificance of evolutionary variation in fin design.     

Volumetric imaging of fish locomotion

Fishes use multiple flexible fins in order to move and maintain stability in a complex fluid environment. We used a new approach, a volumetric velocimetry imaging system, to provide the first instantaneous three-dimensional views of wake structures as they are produced by freely swimming fishes. This new technology allowed us to demonstrate conclusively the linked ring vortex wake pattern that is produced by the symmetrical (homocercal) tail of fishes, and to visualize for the first time the three-dimensional vortex wake interaction between the dorsal and anal fins and the tail. We found that the dorsal and anal fin wakes were rapidly (within one tail beat) assimilated into the caudal fin vortex wake. These results show that volumetric imaging of biologically generated flow patterns can reveal new features of locomotor dynamics, and provides an avenue for future investigations of the diversity of fish swimming patterns and their hydrodynamic consequences.     

Musculoskeletal morphology and regionalization within the dorsal and anal fins of bluegill sunfish (Lepomis macrochirus)

Ray‐finned fishes actively control the shape and orientation of their fins to either generate or resist hydrodynamic forces. Because of the emergent mechanical properties of their segmented, bilaminar fin rays (lepidotrichia), and actuation by multiple muscles, fish can control the rigidity and curvature of individual rays independently, thereby varying the resultant forces across the fin surfaces. Expecting that differences in fin‐ray morphology should reflect variation in their mechanical properties, we measured several musculoskeletal features of individual spines and rays of the dorsal and anal fins of bluegill sunfish, Lepomis macrochirus, and assessed their mobility and flexibility. We separated the fin‐rays into four groups based on the fin (dorsal or anal) or fin‐ray type (spine or ray) and measured the length of the spines/rays and the mass of the three median fin‐ray muscles: the inclinators, erectors and depressors. Within the two ray groups, we measured the portion of the rays that were segmented vs. unsegmented and branched vs. unbranched. For the majority of variables tested, we found that variations between fin‐rays within each group were significantly related to position within the fin and these patterns were conserved between the dorsal and anal rays. Based on positional variations in fin‐ray and muscle parameters, we suggest that anterior and posterior regions of each fin perform different functions when interacting with the surrounding fluid. Specifically, we suggest that the stiffer anterior rays of the soft dorsal and anal fins maintain stability and keep the flow across the fins steady. The posterior rays, which are more flexible with a greater range of motion, fine‐tune their stiffness and orientation, directing the resultant flow to generate lateral and some thrust forces, thus acting as an accessory caudal fin. J. Morphol., 2012. © 2011 Wiley Periodicals, Inc.     

Biomechanics of swimming in the pufferfish Diodon holocanthus: propulsive momentum enhancement is an adaptation for thrust production in an undulatory median and paired-fin swimmer

A form of large-amplitude elongated-body theory appropriate for the analysis of undulatory fins attached to a rigid body of elliptical section suggests a benefit due to momentum enhancement relative to the fins on their own. This theoretical prediction is experimentally confirmed for the first time. Theoretical momentum enhancement factors for Diodon holocanthus (2.2 and 2.7 for the median and pectoral fins, respectively) compared well to inferred thrust values determined from particle-image velocimetry (PIV) wake measurements (2.2-2.4 and 2.7-2.9). Caudal fin mean theoretical thrust was not significantly different from measured (PIV) values (n = 24, P > 0.05), implying no momentum enhancement. Pectoral-fin thrust was half that of the median and caudal fins due to high fin-jet angles, low circulation and momentum. Average total fin thrust and fish drag were not significantly different (n = 24, P > 0.05). Vortex rings generated by the fins were elliptical, with size dependent on fin chord and stroke amplitude. Hydrodynamic advantages (thrust enhancement at no cost to hydrodynamic efficiency, reduction of side forces minimizing energy wasting yawing motions and body drag) are probably common among rigid-bodied organisms propelled by undulatory fins. A trade-off between momentum enhancement and the rate of momentum generation (thrust force) sets a practical limit to the former. For small fins whilst momentum enhancement is high, absolute thrust is low. In addition, previously suggested limitations on thrust enhancement set by reductions in propulsive force associated with progressive reductions in fin wavelength are found to be biologically unrealistic.     

Functional dorsoventral symmetry in relation to lift-based swimming in the ocean sunfish Mola mola

The largest (up to 2 tons) and a globally distributed teleost--the ocean sunfish Mola mola--is commonly regarded as a planktonic fish because of its unusual shape including absence of caudal fin. This common view was recently questioned because the horizontal movements of the ocean sunfish tracked by acoustic telemetry were independent of ocean currents. However, direct information regarding their locomotor performance under natural conditions is still lacking. By using multi-sensor tags, we show that sunfish indeed swam continuously with frequent vertical movements at speeds of 0.4-0.7 m s(-1), which is similar to the records of other large fishes such as salmons, marlins, and pelagic sharks. The acceleration data revealed that they stroked their dorsal and anal fins synchronously (dominant frequency, 0.3-0.6 Hz) to generate a lift-based thrust, as penguins do using two symmetrical flippers. Morphological studies of sunfish (mass, 2-959 kg) showed that the dorsal and anal fins had similar external (symmetrical shape and identical area) and internal (identical locomotory muscle mass) features; however, the muscle shape differed markedly. We conclude that ocean sunfish have functional dorsoventral symmetry with regards to the non-homologous dorsal and anal fins that act as a pair of vertical hydrofoils. Although sunfish lack a swimbladder, we found that they are neutrally buoyant independent of depth because of their subcutaneous gelatinous tissue that has low density and is incompressible. Efficient lift-based swimming in conjunction with neutral buoyancy enables sunfish to travel long distances both horizontally and vertically.     

Use of biorobotic models of highly deformable fins for studying the mechanics and control of fin forces in fishes

Bony fish swim with a level of agility that is unmatched in human-developed systems. This is due, in part, to the ability of the fish to carefully control hydrodynamic forces through the active modulation of the fins' kinematics and mechanical properties. To better understand how fish produce and control forces, biorobotic models of the bluegill sunfish's (Lepomis macrochirus) caudal fin and pectoral fins were developed. The designs of these systems were based on detailed analyses of the anatomy, kinematics, and hydrodynamics of the biological fins. The fin models have been used to investigate how fin kinematics and the mechanical properties of the fin-rays influence propulsive forces and to explore kinematic patterns that were inspired by biological motions but that were not explicitly performed by the fish. Results from studies conducted with the fin models indicate that subtle changes to the kinematics and mechanical properties of fin rays can significantly impact the magnitude, direction, and time course of the 3D forces used for propulsion and maneuvers. The magnitude of the force tends to scale with the fin's stiffness, but the direction of the force is not invariant, and this causes disproportional changes in the magnitude of the thrust, lift, and lateral components of force. Results from these studies shed light on the multiple strategies that are available to the fish to modulate fin forces.     

Correlated evolution of body and fin morphology in the cichlid fishes

Body and fin shapes are chief determinants of swimming performance in fishes. Different configurations of body and fin shapes can suit different locomotor specializations. The success of any configuration is dependent upon the hydrodynamic interactions between body and fins. Despite the importance of body–fin interactions for swimming, there are few data indicating whether body and fin configurations evolve in concert, or whether these structures vary independently. The cichlid fishes are a diverse family whose well‐studied phylogenetic relationships make them ideal for the study of macroevolution of ecomorphology. This study measured body, and caudal and median fin morphology from radiographs of 131 cichlid genera, using morphometrics and phylogenetic comparative methods to determine whether these traits exhibit correlated evolution. Partial least squares canonical analysis revealed that body, caudal fin, dorsal fin, and anal fin shapes all exhibited strong correlated evolution consistent with locomotor ecomorphology. Major patterns included the evolution of deep body profiles with long fins, suggestive of maneuvering specialization; and the evolution of narrow, elongate caudal peduncles with concave tails, a combination that characterizes economical cruisers. These results demonstrate that body shape evolution does not occur independently of other traits, but among a suite of other morphological changes that augment locomotor specialization.     

Structure of supporting elements in the dorsal fin of percid fishes

The dorsal fin is one of the most varied swimming structures in Acanthomorpha, the spiny‐finned fishes. This fin can be present as a single contiguous structure supported by bony spines and soft lepidotrichia, or it may be divided into an anterior, spiny dorsal fin and a posterior, soft dorsal fin. The freshwater fish family Percidae exhibits especially great variation in dorsal fin spacing, including fishes with separated fins of varying gap length and fishes with contiguous fins. We hypothesized that fishes with separated dorsal fins, especially those with large gaps between fins, would have stiffened fin elements at the leading edge of the soft dorsal fin to resist hydrodynamic loading during locomotion. For 10 percid species, we measured the spacing between dorsal fins and calculated the second moment of area of selected spines and lepidotrichia from museum specimens. There was no significant relationship between the spacing between dorsal fins and the second moment of area of the leading edge of the soft dorsal fin.     

New genus and species of the family stichaeidae from Hokkaido,Japan

Specimens of a new genus and species of the stichaeid fish,Leptostichaeus pumilus, were collected from the Okhotsk Sea off Hokkaido in Japan. The present new genus and species clearly differs from all the other genera and species of the stichaeid fishes in the following characters: 3 or 4 pectoral fin rays; 10 or fewer caudal principal rays; 79–82 dorsal spines; no pelvic fin; last interneural spine supporting a single dorsal spine; infraorbital, occipital and lateral line canals absent; moderate size of dorsal spine shorter than eye diameter; membranes of dorsal and anal fins widely connected with caudal fin; a large black spot divided by a yellow band present just above gill cover.     

The fine structure of the fin musculature in two teleost species with different swimming modes,the puffer,Tetraodon steindachneri,and the goldfish,Carassius auratus

Summary Puffer fish (Tetraodon steindachneri) can execute precise maneuvers due to their highly specialized mode of propulsion. In the conventional locomotion exemplified by the goldfish (Carassius auratus), the fish thrusts are generated by lateral beating of the caudal fin. In contrast, the puffer generates its propulsive force by very rapid undulating movements of the pectoral, dorsal and anal fins. The fine structure of the fin muscles is identical in the two species of fishes, despite the differences in fin movement; cytologically, the fibers are intermediate between those of red and of white muscle. On the other hand, both the fusion frequency and the number of motor endplates are considerably higher in the fin muscles of the puffer than in those of the goldfish.     

Resolving Shifting Patterns of Muscle Energy Use in Swimming Fish

Muscle metabolism dominates the energy costs of locomotion. Although in vivo measures of muscle strain, activity and force can indicate mechanical function, similar muscle-level measures of energy use are challenging to obtain. Without this information locomotor systems are essentially a black box in terms of the distribution of metabolic energy. Although in situ measurements of muscle metabolism are not practical in multiple muscles, the rate of blood flow to skeletal muscle tissue can be used as a proxy for aerobic metabolism, allowing the cost of particular muscle functions to be estimated. Axial, undulatory swimming is one of the most common modes of vertebrate locomotion. In fish, segmented myotomal muscles are the primary power source, driving undulations of the body axis that transfer momentum to the water. Multiple fins and the associated fin muscles also contribute to thrust production, and stabilization and control of the swimming trajectory. We have used blood flow tracers in swimming rainbow trout (Oncorhynchus mykiss) to estimate the regional distribution of energy use across the myotomal and fin muscle groups to reveal the functional distribution of metabolic energy use within a swimming animal for the first time. Energy use by the myotomal muscle increased with speed to meet thrust requirements, particularly in posterior myotomes where muscle power outputs are greatest. At low speeds, there was high fin muscle energy use, consistent with active stability control. As speed increased, and fins were adducted, overall fin muscle energy use declined, except in the caudal fin muscles where active fin stiffening is required to maintain power transfer to the wake. The present data were obtained under steady-state conditions which rarely apply in natural, physical environments. This approach also has potential to reveal the mechanical factors that underlie changes in locomotor cost associated with movement through unsteady flow regimes.     

Mapping QTL for an Adaptive Trait: The Length of Caudal Fin in <Emphasis Type="Italic">Lates calcarifer</Emphasis>

The caudal fin represents a fundamental design feature of fishes and plays an important role in locomotor dynamics in fishes. The shape of caudal is an important parameter in traditional systematics. However, little is known about genes involved in the development of different forms of caudal fins. This study was conducted to identify and map quantitative trait loci (QTL) affecting the length of caudal fin and the ratio between tail length and standard body length in Asian seabass (Lates calcarifer). One F1 family containing 380 offspring was generated by crossing two unrelated individuals. One hundred and seventeen microsatellites almost evenly distributed along the whole genome were genotyped. Length of caudal fin at 90 days post-hatch was measured. QTL analysis detected six significant (genome-wide significant) and two suggestive (linkage-group-wide significant) QTL on seven linkage groups. The six significant QTL explained 5.5–16.6% of the phenotypic variance, suggesting these traits were controlled by multiple genes. Comparative genomics analysis identified several potential candidate genes for the length of caudal fin. The QTL for the length of caudal fin detected for the first time in marine fish may provide a starting point for the future identification of genes involved in the development of different forms of caudal fins in fishes.     

Effect of an Artificial Caudal Fin on the Performance of a Biomimetic Fish Robot Propelled by Piezoelectric Actuators

This paper addresses the design of a biomimetic fish robot actuated by piezoeeramic actuators and the effect of artificial caudal fins on the fish robot＇s performance. The limited bending displacement produced by a lightweight piezocomposite actuator was amplified and transformed into a large tail beat motion by means of a linkage system. Caudal fins that mimic the shape of a mackerel fin were fabricated for the purpose of examining the effect of caudal fm characteristics on thrust production at an operating frequency range. The thickness distribution of a real mackerel＇s fin was measured and used to design artificial caudal fins. The thrust performance of the biomimetic fish robot propelled by fins of various thicknesses was examined in terms of the Strouhal number, the Froude number, the Reynolds number, and the power consumption. For the same fm area and aspect ratio, an artificial caudal fin with a distributed thickness shows the best forward speed and the least power consumption.     

Disentangling the functional roles of morphology and motion in the swimming of fish

In fishes the shape of the body and the swimming mode generally are correlated. Slender-bodied fishes such as eels, lampreys, and many sharks tend to swim in the anguilliform mode, in which much of the body undulates at high amplitude. Fishes with broad tails and a narrow caudal peduncle, in contrast, tend to swim in the carangiform mode, in which the tail undulates at high amplitude. Such fishes also tend to have different wake structures. Carangiform swimmers generally produce two staggered vortices per tail beat and a strong downstream jet, while anguilliform swimmers produce a more complex wake, containing at least two pairs of vortices per tail beat and relatively little downstream flow. Are these differences a result of the different swimming modes or of the different body shapes, or both? Disentangling the functional roles requires a multipronged approach, using experiments on live fishes as well as computational simulations and physical models. We present experimental results from swimming eels (anguilliform), bluegill sunfish (carangiform), and rainbow trout (subcarangiform) that demonstrate differences in the wakes and in swimming performance. The swimming of mackerel and lamprey was also simulated computationally with realistic body shapes and both swimming modes: the normal carangiform mackerel and anguilliform lamprey, then an anguilliform mackerel and carangiform lamprey. The gross structure of simulated wakes (single versus double vortex row) depended strongly on Strouhal number, while body shape influenced the complexity of the vortex row, and the swimming mode had the weakest effect. Performance was affected even by small differences in the wakes: both experimental and computational results indicate that anguilliform swimmers are more efficient at lower swimming speeds, while carangiform swimmers are more efficient at high speed. At high Reynolds number, the lamprey-shaped swimmer produced a more complex wake than the mackerel-shaped swimmer, similar to the experimental results. Finally, we show results from a simple physical model of a flapping fin, using fins of different flexural stiffness. When actuated in the same way, fins of different stiffnesses propel themselves at different speeds with different kinematics. Future experimental and computational work will need to consider the mechanisms underlying production of the anguilliform and carangiform swimming modes, because anguilliform swimmers tend to be less stiff, in general, than are carangiform swimmers.     

Morphological scaling of body form in four shark species differing in ecology and life history

Body form can change across ontogeny, and can influence how animals of different sizes move and feed. Scaling data on live apex predatory sharks are rare and, therefore, we examined patterns of scaling in ontogenetic series of four sympatric shark species exhibiting a range of sizes, ecologies and life histories (tiger, bull, blacktip, and nurse shark). We evaluated 13 linear morphological variables and two areas (caudal and dorsal) that could influence both animal condition and locomotor performance. These measurements included dimensions of the dorsal, pectoral, and caudal fins, as well as several dimensions of body circumference, and of the head. For all four species, the body axis (eye‐to‐eye, lateral span, frontal span, proximal span) scaled close to isometry (expected slope of 1.0). The two largest sharks (tiger and bull sharks) also showed significant negative allometry for elements of the caudal fin. We found significant negative allometry in the lengths of the upper lobe of the caudal fin (caudal fin 1) and the overall height of the caudal fin (caudal fin 2) in tiger and bull sharks, with slopes ranging from about 0.60 to 0.73. Further, tiger sharks showed negative allometry in caudal fin area. These results suggest that in terms of overall body dimensions, small sharks are roughly geometrically similar to large sharks, at least within the species we examined. However, juvenile tiger (and to a lesser extent bull sharks) are notable in having proportionately larger caudal fins compared to adult sharks. As the caudal fin contributes to generating thrust during forward locomotion, this scaling implies differences among adult and juvenile sharks in locomotor ability. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 126–135.     

Swimming in four goldfish Carassius auratus morphotypes: understanding functional design and performance employing artificially selected forms

Four goldfish Carassius auratus morphotypes of similar length (50 mm): common (streamlined, full complement of paired and median fins, bifurcated caudal fin), comet (‘common like’ but with a long, deeply forked caudal fin), fantail (short, deep body with twinned caudal and anal fins) and eggfish (similar to the fantail but lacking a dorsal fin) were compared. Drag, steady swimming kinematics, energetics, fast‐start performance, stability in yaw and roll and propulsive muscle ultrastructural characteristics were measured. A performance ‘pairing’ (common and comet; fantail and eggfish) was a recurrent theme for most performance variables. Fantail and eggfish drag were higher (requiring more thrust at any given velocity) than those for the more streamlined common and comet. This was reflected in kinematics; tailbeat frequency and stride length at any given velocity for the common and comet were lower and higher, respectively, than that of the fantail and eggfish. Common and comet fatigue times were not significantly different from those of their ancestor, crucian carp Carassius carassius, and higher than the fantail and eggfish. The cost of transport of the common and comet (c. 0· 6 mg O₂ kg^?1 m^?1) was accurately predicted by the mass scaling relationship for fish, but values for the fantail and eggfish (c. 1· 3 mg O₂ kg^?1 m^?1) were not. Rolling and yawing motions in eggfish (dorsal fin absent) during steady swimming were associated with significant energy losses. Eggfish maximum fast‐start acceleration (c. 5 m s^?2) was poor due to the absence of inertial and lifting contributions to thrust from the dorsal fin and energy wasting rolling motions. Common and comet fast‐start performance (average velocity c. 0· 45 m s^?1, maximum velocity c. 1· 2 m s^?1, average acceleration c. 7· 5 m s^?2, maximum acceleration c. 35 m s^?2) was similar to that of other locomotor generalists (e.g. rainbow trout Oncorhynchus mykiss). Artificially selected fishes can contribute to the understanding of form and movement in fishes and complement studies of the role of locomotor adaptations in natural systems.     

Mechanosensation in an adipose fin

Adipose fins are found on approximately 20% of ray-finned fish species. The apparently rudimentary anatomy of adipose fins inspired a longstanding hypothesis that these fins are vestigial and lack function. However, adipose fins have evolved repeatedly within Teleostei, suggesting adaptive function. Recently, adipose fins were proposed to function as mechanosensors, detecting fluid flow anterior to the caudal fin. Here we test the hypothesis that adipose fins are mechanosensitive in the catfish Corydoras aeneus. Neural activity, recorded from nerves that innervate the fin, was shown to encode information on both movement and position of the fin membrane, including the magnitude of fin membrane displacement. Thus, the adipose fin of C. aeneus is mechanosensitive and has the capacity to function as a ‘precaudal flow sensor’. These data force re-evaluation of adipose fin clipping, a common strategy for tagging fishes, and inform hypotheses of how function evolves in novel vertebrate appendages.     

Function of dorsal fins in bamboo shark during steady swimming

To gain insight into the function of the dorsal fins in white-spotted bamboo sharks (Orectolobiformes: Hemiscyillidae) during steady swimming, data on three-dimensional kinematics and electromyographic recordings were collected. Bamboo sharks were induced to swim at 0.5 and 0.75 body lengths per second in a laminar flow tank. Displacement, lag and angles were analyzed from high-speed video images. Onset, offset, duration, duty cycle and asynchrony index were calculated from three muscle implants on each side of each dorsal fin. The dorsal fins were displaced more laterally than the undulating body. In addition, the dorsal tips had larger lateral displacement than the trailing edges. Increased speed was accompanied by an increase in tail beat frequency with constant tail beat amplitude. However, lateral displacement of the fins and duration of muscle bursts remained relatively constant with increased speed. The range of lateral motion was greater for the second dorsal fin (mean 33.3°) than for the first dorsal fin (mean 28.4°). Bending within the fin was greater for the second dorsal fin (mean 43.8°) than for the first dorsal fin (mean 30.8°). Muscle onset and offset among implants on the same side of each dorsal fin was similar. Three-dimensional conformation of the dorsal fins was caused by interactions between muscle activity, material properties, and incident flow. Alternating bilateral activity occurred in both dorsal fins, further supporting the active role of these hydrofoils in thrust production during steady swimming. The dorsal fins in bamboo sharks are capable of thrust production during steady swimming and do not appear to function as stabilizing structures.     

Design and Implementation of Paired Pectoral Fins Locomotion of Labriform Fish Applied to a Fish Robot

In present,there are increasing interests in the research on mechanical and control system of underwater vehicles.Theseongoing research efforts are motivated by more pervasive applications of such vehicles including seabed oil and gas explorations,scientific deep ocean surveys,military purposes,ecological and water environmental studies,and also entertainments.However,the performance of underwater vehicles with screw type propellers is not prospective in terms of its efficiency andmaneuverability.The main weaknesses of this kind of propellers are the production of vortices and sudden generation of thrustforces which make the control of the position and motion difficult.On the other hand,fishes and other aquatic animals are efficient swimmers,posses high maneuverability,are able to followtrajectories,can efficiently stabilize themselves in currents and surges,create less wakes than currently used underwater vehicle,and also have a noiseless propulsion.The fish’s locomotion mechanism is mainly controlled by its caudal fin and paired pectoralfins.They are classified into Body and/or Caudal Fin(BCF)and Median and/or paired Pectoral Fins(MPF).The study of highlyefficient swimming mechanisms of fish can inspire a better underwater vehicles thruster design and its mechanism.There are few studies on underwater vehicles or fish robots using paired pectoral fins as thruster.The work presented in thispaper represents a contribution in this area covering study,design and implementation of locomotion mechanisms of pairedpectoral fins in a fish robot.The performance and viability of the biomimetic method for underwater vehicles are highlightedthrough in-water experiment of a robotic fish.