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Swimming in schools has long been hypothesized to allow fish to save energy. Fish must exploit the energy from the wakes of their neighbors for maximum energy savings, a feat that requires them to both synchronize their tail movements and stay in certain positions relative to their neighbors. To maintain position in a school, we know that fish use multiple sensory systems, mainly their visual and flow sensing lateral line system. However, how fish synchronize their swimming movements in a school is still not well understood. Here, we test the hypothesis that this synchronization may depend on functional differences in the two branches of the lateral line sensory system that detects water movements close to the fish's body. The anterior branch, located on the head, encounters largely undisturbed free-stream flow, while the posterior branch, located on the trunk and tail, encounters flow that has been affected strongly by the tail movement. Thus, we hypothesize that the anterior branch may be more important for regulating position within the school, while the posterior branch may be more important for synchronizing tail movements. Our study examines functional differences in the anterior and posterior lateral line in the structure and tail synchronization of fish schools. We used a widely available aquarium fish that schools, the giant danio, Devario equipinnatus. Fish swam in a large circular tank where stereoscopic videos recordings were used to reconstruct the 3D position of each individual within the school and to track tail kinematics to quantify synchronization. For one fish in each school, we ablated using cobalt chloride either the anterior region only, the posterior region only, or the entire lateral line system. We observed that ablating any region of the lateral line system causes fish to swim in a "box" or parallel swimming formation, which was different from the diamond formation observed in normal fish. Ablating only the anterior region did not substantially reduce tail beat synchronization but ablating only the posterior region caused fish to stop synchronizing their tail beats, largely because the tail beat frequency increased dramatically. Thus, the anterior and posterior lateral line system appears to have different behavioral functions in fish. Most importantly, we showed that the posterior lateral line system played a major role in determining tail beat synchrony in schooling fish. Without synchronization, swimming efficiency decreases, which can have an impact on the fitness of the individual fish and group. [ABSTRACT FROM AUTHOR]

The anterior body of many fishes is shaped like an airfoil turned on its side. With an oscillating angle to the swimming direction, such an airfoil experiences negative pressure due to both its shape and pitching movements. This negative pressure acts as thrust forces on the anterior body. Here, we apply a high-resolution, pressurebased approach to describe how two fishes, bluegill sunfish (Lepomis macrochirus Rafinesque) and brook trout (Salvelinus fontinalis Mitchill), swimming in the carangiform mode, the most common fish swimming mode, generate thrust on their anterior bodies using leading-edge suction mechanics, much like an airfoil. These mechanics contrast with those previously reported in lampreys— anguilliform swimmers—which produce thrust with negative pressure but do so through undulatory mechanics. The thrust produced on the anterior bodies of these carangiform swimmers through negative pressure comprises 28% of the total thrust produced over the body and caudal fin, substantially decreasing the net drag on the anterior body. On the posterior region, subtle differences in body shape and kinematics allow trout to produce more thrust than bluegill, suggesting that they may swim more effectively. Despite the large phylogenetic distance between these species, and differences near the tail, the pressure profiles around the anterior body are similar. We suggest that such airfoil-like mechanics are highly efficient, because they require very little movement and therefore relatively little active muscular energy, and may be used by a wide range of fishes since many species have appropriately shaped bodies. [ABSTRACT FROM AUTHOR]

Fish live in a complex world and must actively adapt their swimming behavior to a range of environments. Most studies of swimming kinematics focus on two-dimensional properties related to the bending wave that passes from head to tail. However, fish also twist their bodies three dimensionally around their longitudinal axis as the bending wave passes down the body. We measured and characterized this movement, which we call 'wobble', in six species of elongate fishes (Anoplarchus insignis, Xiphister mucosus, Lumpenus sagitta, Pholis laeta, Apodichthys flavidus and Ronquilus jordani) from three different habitats (intertidal, nearshore and subtidal) using custom video analysis software. Wobble and bending are synchronized, with a phase shift between the wobble wave and bending wave. We found that species from the same habitats swim in similar ways, even if they are more closely related to species from different habitats. In nearshore species, the tail wobbles the most but, in subtidal and intertidal species, the head wobbles more than or the same as the tail. We also wanted to understand the relationship between wobble and the passive mechanics of the fish bodies. Therefore, we measured torsional stiffness and modulus along the body and found that modulus increases from head to tail in all six species. As wobble does not correlate with the passive properties of the body, it may play a different role in swimming behavior of fishes from different habitats. [ABSTRACT FROM AUTHOR]

In their natural environment, fish must swim stably through unsteady flows and vortices, including vertical vortices, typically shed by posts in a flow, horizontal cross-flow vortices, often produced by a step or a waterfall in a stream, and streamwise vortices, where the axis of rotation is aligned with the direction of the flow. Streamwise vortices are commonly shed by bluff bodies in streams and by ships' propellers and axial turbines, but we know little about their effects on fish. Here, we describe how bluegill sunfish use more energy and are destabilized more often in flow with strong streamwise vorticity. The vortices were created inside a sealed flow tank by an array of four turbines with similar diameter to the experimental fish. We measured oxygen consumption for seven sunfish swimming at 1.5 body lengths (BL) s~1 with the turbines rotating at 2 Hz and with the turbines off (control). Simultaneously, we filmed the fish ventrally and recorded the fraction of time spent maneuvering side-to-side and accelerating forward. Separately, we also recorded lateral and ventral video for a combination of swimming speeds (0.5, 1.5 and 2.5 BL s-1) and turbine speeds (0, 1, 2 and 3 Hz), immediately after turning the turbines on and 10 min later to test for accommodation. Bluegill sunfish are negatively affected by streamwise vorticity. Spills (loss of heading), maneuvers and accelerations were more frequent when the turbines were on than in the control treatment. These unsteady behaviors, particularly acceleration, correlated with an increase in oxygen consumption in the vortex flow. Bluegill sunfish are generally fast to recover from roll perturbations and do so by moving their pectoral fins. The frequency of spills decreased after the turbines had run for 10 min, but was still markedly higher than in the control, showing that fish partially adapt to streamwise vorticity, but not completely. Coping with streamwise vorticity may be an important energetic cost for stream fishes or migratory fishes. [ABSTRACT FROM AUTHOR]

Abstract: The bodies of many fishes are flexible, elastic structures; if you bend them, they spring back. Therefore, they should have a resonant frequency: a bending frequency at which the output amplitude is maximized for a particular input. Previous groups have hypothesized that swimming at this resonant frequency could maximize efficiency, and that a neural circuit called the central pattern generator might be able to entrain to a mechanical resonance. However, fishes swim in water, which may potentially damp out many resonant effects. Additionally, their bodies are elongated, which means that bending can occur in complicated ways along the length of the body. We review previous studies of the mechanical properties of fish bodies, and then present new data that demonstrate complex bending properties of elongated fish bodies. Resonant peaks in amplitude exist, but there may be many of them depending on the body wavelength. Additionally, they may not correspond to the maximum swimming speed. Next, we describe experiments using a closed-loop preparation of the lamprey, in which a preparation of the spinal cord is linked to a real-time simulation of the muscle and body properties, allowing us to examine resonance entrainment as we vary the simulated resonant frequency. We find that resonance entrainment does occur, but is rare. Gain had a significant, though weak, effect, and a nonlinear muscle model produced resonance entrainment more often than a linear filter. We speculate that resonance may not be a critical effect for efficient swimming in elongate, anguilliform swimmers, though it may be more important for stiffer carangiform and thunniform fishes. [Copyright &y& Elsevier]

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. [ABSTRACT FROM AUTHOR]

Fishes have an enormous diversity of body shapes and fin morphologies. From a hydrodynamic standpoint, the functional significance of this diversity is poorly understood, largely because the three-dimensional flow around swimming fish is almost completely unknown. Fully three-dimensional volumetric flow measurements are not currently feasible, but measurements in multiple transverse planes along the body can illuminate many of the important flow features. In this study, I analyze flow in the transverse plane at a range of positions around bluegill sunfish Lepomis macrochirus, from the trailing edges of the dorsal and anal fins to the near wake. Simultaneous particle image velocimetry and kinematic measurements were performed during swimming at 1.2 body lengths s-1 to describe the streamwise vortex structure, to quantify the contributions of each fin to the vortex wake, and to assess the importance of three-dimensional flow effects in swimming. Sunfish produce streamwise vortices from at least eight distinct places, including both the dorsal and ventral margins of the soft dorsal and anal fins, and the tips and central notched region of the caudal fin. I propose a three-dimensional structure of the vortex wake in which these vortices from the caudal notch are elongated by the dorso-ventral cupping motion of the tail, producing a structure like a hairpin vortex in the caudal fin vortex ring. Vortices from the dorsal and anal fin persist into the wake, probably linking up with the caudal fin vortices. These dorsal and anal fin vortices do not differ significantly in circulation from the two caudal fin tip vortices. Because the circulations are equal and the length of the trailing edge of the caudal fin is approximately equal to the combined trailing edge length of the dorsal and anal fins, I argue that the two anterior median fins produce a total force that is comparable to that of the caudal fin. To provide additional detail on how different positions contribute to total force along the posterior body, the change in vortex circulation as flow passes down the body is also analyzed. The posterior half of the caudal fin and the dorsal and anal fins add vortex circulation to the flow, but circulation appears to decrease around the peduncle and anterior caudal fin. Kinematic measurements indicate that the tail is angled correctly to enhance thrust through this interaction. Finally, the degree to which the caudal fin acts like a idealized two-dimensional plate is examined: approximately 25% of the flow near the tail is accelerated up and down, rather than laterally, producing wasted momentum, a loss not present in ideal two-dimensional theories. [ABSTRACT FROM AUTHOR]