Person:

Lauder, George

There are approximately 50 muscles that control tail fin shape in most teleost fishes, and although myotomal muscle function has been extensively studied, little work has been done on the intrinsic musculature that controls and shapes the tail. In this study we measured electrical activity in intrinsic tail musculature to determine if these muscles are active during steady rectilinear locomotion, and to compare intrinsic muscle recruitment patterns to previous data on myotomal muscle fibers. Five bluegill sunfish (Lepomis macrochirus) were anaesthetized and electrode wires surgically placed into a total of 24 intrinsic caudal muscles, up to 13 at a time, and activity was correlated with synchronous recordings from myotomal fibers in the caudal peduncle. After recovery, fish swam steadily at speeds of 0.5, 1.2 and 2.0·L·s–1, while filmed from lateral, posterior and ventral views simultaneously at 250·frames·s–1. Comparison among speeds confirmed that muscle recruitment varies significantly with speed. At 0.5·L·s–1, the caudal fin was generally not used for propulsion, and swimming was accomplished primarily through body undulations. Intrinsic caudal muscle activity at this speed was intermittent and variable. At 1.2 and 2.0·L·s–1, the supracarinalis and infracarinalis muscles acted on the dorsal- and ventral-most fin rays, respectively, to expand the surface area of the caudal fin. The interradialis muscles adducted individual fin rays, dorsally to ventrally, following activation of the hypochordal longitudinalis. Contralateral muscle activity of interradialis muscles occurred as the caudal fin crossed the mean direction of travel and fin height was greatest, whereas ipsilateral activity of carinalis muscles occurred near points of maximum excursion of the fin, at speeds of 1.2 and 2.0·L·s–1, after fin height was lowest. Burst intensity increased with swimming speed, suggesting stiffening of the tail fin against imposed hydrodynamic loads. Activity patterns of intrinsic caudal muscles suggest that these most posterior muscles in fishes, located within the tail, are among the very first recruited as swimming speed increases, and that slow undulatory swimming is powered by muscle fibers located posteriorly in the caudal peduncle and tail.

As a result of years of research on the comparative biomechanics and physiology of moving through water, biologists and engineers have made considerable progress in understanding how animals moving underwater use their muscles to power movement, in describing body and appendage motion during propulsion, and in conducting experimental and computational analyses of fluid movement and attendant forces. But it is clear that substantial future progress in understanding aquatic propulsion will require new lines of attack. Recent years have seen the advent of one such new avenue that promises to greatly broaden the scope of intellectual opportunity available to researchers: the use of biorobotic models. In this paper we discuss, using aquatic propulsion in fishes as our focal example, how using robotic models can lead to new insights in the study of aquatic propulsion. We use two examples: (1) pectoral fin function, and (2) hydrodynamic interactions between dorsal and caudal fins. Pectoral fin function is characterized by considerable deformation of individual fin rays, as well as spanwise (along the length) and chordwise (across the fin) deformation and area change. The pectoral fin can generate thrust on both the outstroke and instroke. A robotic model of the pectoral fin replicates this result, and demonstrates the effect of altering stroke kinematics on the pattern of force production. The soft dorsal fin of fishes sheds a distinct vortex wake that dramatically alters incoming flow to the tail: the dorsal fin and caudal fin act as dual flapping foils in series. This design can be replicated with a dual-foil flapping robotic device that demonstrates this phenomenon and allows examination of regions of the flapping performance space not available to fishes. We show how the robotic flapping foil device can also be used to better understand the significance of flexible propulsive surfaces for locomotor performance. Finally we emphasize the utility of self-propelled robotic devices as a means of understanding how locomotor forces are generated, and review different conceptual designs for robotic models of aquatic propulsion.

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.