Person:

Lauder, George

The fast-start escape response is critically important to avoid predation, and axial movements driving it have been studied intensively. Large median dorsal and anal fins located near the tail have been hypothesized to increase acceleration away from the threat, yet the contribution of flexible median fins remains undescribed. To investigate the role of median fins, C-start escape responses of bluegill sunfish (Lepomis macrochirus) were recorded by three high-speed, high-resolution cameras at 500framess–1 and the 3-D kinematics of individual dorsal and anal fin rays were analyzed. Movement and orientation of the fin rays relative to the body axis were calculated throughout the duration of the C-start. We found that: (1) timing and magnitude of angular displacement varied among fin rays based on position within the fin and (2) kinematic patterns support the prediction that fin rays are actively resisting hydrodynamic forces and transmitting momentum into the water. We suggest that regions within the fins have different roles. Anterior regions of the fins are rapidly elevated to increase the volume of water that the fish may interact with and transmit force into, thus generating greater total momentum. The movement pattern of all the fin rays creates traveling waves that move posteriorly along the length of the fin, moving water as they do so. Flexible posterior regions ultimately act to accelerate this water towards the tail, potentially interacting with vortices generated by the caudal fin during the C-start. Despite their simple appearance, median fins are highly complex and versatile control surfaces that modulate locomotor performance.

Although kinematic analysis of individual fin rays provides valuable insight into the contribution of median fins to C-start performance, it paints an incomplete picture of the complex movements and deformation of the flexible fin surface. To expand our analysis of median fin function during the escape response of bluegill sunfish (Lepomis macrochirus), patterns of spanwise and chordwise curvature of the soft dorsal and anal fin surfaces were examined from the same video sequences previously used in analysis of fin-ray movement and orientation. We found that both the span and chord undergo undulation, starting in the anterior region of either fin. Initiated early in Stage 1 of the C-start, the undulation travels in a postero-distal direction, reaching the trailing edge of the fins during early Stage 2. Maximum spanwise curvature typically occurred among the more flexible posterior fin rays, though there was no consistent correlation between maximum curvature and fin-ray position. Undulatory patterns suggest different mechanisms of action for the fin regions. In the anterior fin region, where the fin rays are oriented dorsoventrally, undulation is directed primarily chordwise, initiating a transfer of momentum into the water to overcome the inertia of the flow and direct the water posteriorly. Within the posterior region, where the fin rays are oriented caudally, undulation is predominantly directed spanwise; thus, the posterior fin region acts to ultimately accelerate this water towards the tail to increase thrust forces. Treatment of median fins as appendages with uniform properties does not do justice to their complexity and effectiveness as control surfaces.

Recent kinematic and hydrodynamic studies on fish median fins have shown that dorsal fins actively produce jets with large lateral forces. Because of the location of dorsal fins above the fish’s rolling axis, these lateral forces, if unchecked, would cause fish to roll. In this paper we examine the hydrodynamics of trout anal fin function and hypothesize that anal fins, located below the fish’s rolling axis, produce similar jets to the dorsal fin and help balance rolling torques during swimming. We simultaneously quantify the wake generated by dorsal and anal fins in brook trout by swimming fish in two horizontal light sheets filmed by two synchronized high speed cameras during steady swimming and manoeuvring. Six major conclusions emerge from these experiments. First, anal fins produce lateral jets to the same side as dorsal fins, confirming the hypothesis that anal fins produce fluid jets that balance those produced by dorsal fins. Second, in contrast to previous work on sunfish, neither dorsal nor anal fins produce significant thrust during steady swimming; flow leaves the dorsal and anal fins in the form of a shear layer that rolls up into vortices similar to those seen in steady swimming of eels. Third, dorsal and anal fin lateral jets are more coincident in time than would be predicted from simple kinematic expectations; shape, heave and pitch differences between fins, and incident flow conditions may account for the differences in timing of jet shedding. Fourth, relative force and torque magnitudes of the anal fin are larger than those of the dorsal fin; force differences may be due primarily to a larger span and a more squarely shaped trailing edge of the anal fin compared to the dorsal fin; torque differences are also strongly influenced by the location of each fin relative to the fish’s centre of mass. Fifth, flow is actively modified by dorsal and anal fins resulting in complex flow patterns surrounding the caudal fin. The caudal fin does not encounter free-stream flow, but rather moves through incident flow greatly altered by the action of dorsal and anal fins. Sixth, trout anal fin function differs from dorsal fin function; although dorsal and anal fins appear to cooperate functionally, there are complex interactions between other fins and free stream perturbations that require independent dorsal and anal fin motion and torque production to maintain control of body position.

Fish swimming has often been simplified into the motions of a two-dimensional slice through the horizontal midline, as though fishes live in a flat world devoid of a third dimension. While fish bodies do undulate primarily horizontally, this motion has important three-dimensional components, and fish fins can move in a complex three-dimensional manner. Recent results suggest that an understanding of the three-dimensional body shape and fin motions is vital for explaining the mechanics of swimming, and that two-dimensional representations of fish locomotion are misleading. In this study, we first examine axial swimming from the twodimensional viewpoint, detailing the limitations of this view. Then we present data on the kinematics and hydrodynamics of the dorsal fin, the anal fin and the caudal fin during steady swimming and maneuvering in brook trout, Salvelinus fontinalis, bluegill sunfish, Lepomis macrochirus, and yellow perch, Perca flavescens. These fishes actively move the dorsal and anal fins during swimming, resulting in curvature along both anterio-posterior and dorso-ventral axes. The momentum imparted to the fluid by these fins comprises a substantial portion of total swimming force, adding to thrust and contributing to roll stability. While swimming, the caudal fin also actively curves dorso-ventrally, producing vortices separately from both its upper and lower lobes. This functional separation of the lobes may allow additional control of three-dimensional orientation, but probably reduces swimming efficiency. In contrast, fish may boost the caudal finʼs efficiency by taking advantage of the flow from the dorsal and anal fins as it interacts with the flow around the caudal fin itself. During maneuvering, fish readily use their fins outside of the normal planes of motion. For example, the dorsal fin can flick laterally, orienting its surface perpendicular to the body, to help in turning and braking. These data demonstrate that, while fish do move primarily in the horizontal plane, neither their bodies nor their motions can accurately be simplified in a two-dimensional representation. To begin to appreciate the functional consequences of the diversity of fish body shapes and locomotor strategies, one must escape Flatland to examine all three dimensions.

Dorsal and anal fins are median fins located above and below the centre of mass of fishes, each having a moment arm relative to the longitudinal axis. Understanding the kinematics of dorsal and anal fins may elucidate how these fins are used in concert to maintain and change fish body position and yet little is known about the functions of these fins. Using three synchronized high-speed cameras (500·frames·s–1) we studied the three-dimensional kinematics of dorsal and anal fins during steady swimming (0.5–2.5·TL·s–1, where TL=total length) and during slow speed maneuvers (0.5·TL·s–1). By digitizing points along every other fin ray in the soft-rayed portion of the fins we were able to determine not only the movement of the fin surface but also the curvature of individual fin rays and the resulting fin surface shape. We found that dorsal and anal fins begin oscillating, in phase, at steady swimming speeds above 1.0·TL·s–1 and that maximum lateral displacement of the trailing edge of the fins as well as fin area increase with increasing steady swimming speed. Differences in area, lateral displacement and moment arm between the dorsal and anal fin suggest that dorsal and anal fins produce balancing torques during steady swimming. During maneuvers, fin area is maximized and mean lateral excursion of both fins is greater than during steady swimming, with large variation among maneuvers. Fin surface shape changes dramatically during maneuvers. At any given point in time the spanwise (base to tip) curvature along fin rays can differ between adjacent rays, suggesting that fish have a high level of control over fin surface shape. Also, during maneuvers the whole surface of both dorsal and anal fins can be bent without individual fin rays exhibiting significant curvature.