Person:

Lauder, George

Evolutionary patterns of intrinsic caudal musculature in ray-finned fishes show that fine control of the dorsal lobe of the tail evolved first, followed by the ability to control the ventral lobe. This progression of increasing differentiation of musculature suggests specialization of caudal muscle roles. Fine control of fin elements is probably responsible for the range of fin conformations observed during different maneuvering behaviors. Here, we examine the kinematics of the caudal fin and the motor activity of the intrinsic caudal musculature during kick-and-glide, braking and backing maneuvers, and compare these data with our previous work on the function of the caudal fin during steady swimming. Kick-and-glide maneuvers consisted of largeamplitude, rapid lateral excursion of the tail fin, followed by forward movement of the fish with the caudal fin rays adducted to reduce surface area and with the tail held in line with the body. Just before the kick, the flexors dorsalis and ventralis, hypochordal longitudinalis, infracarinalis and supracarinalis showed strong activity. During braking, the dorsal and ventral lobes of the tail moved in opposite directions, forming an ʻSʼ-shape, accompanied by strong activity in the interradialis muscles. During backing up, the ventral lobe initiated a dorsally directed wave along the distal edge of the caudal fin. The relative timing of the intrinsic caudal muscles varied between maneuvers, and their activation was independent of the activity of the red muscle of the axial myomeres in the caudal region. There was no coupling of muscle activity duration and electromyographic burst intensity in the intrinsic caudal muscles during maneuvers, as was observed in previous work on steady swimming. Principal-component analysis produced four components that cumulatively explained 73.6% of the variance and segregated kick-and-glide, braking and backing maneuvers from each other and from steady swimming. The activity patterns of the intrinsic caudal muscles during maneuvering suggest motor control independent from myotomal musculature, and specialization of individual muscles for specific kinematic roles.

Eels undulate a larger portion of their bodies while swimming than many other fishes, but the hydrodynamic consequences of this swimming mode are poorly understood. In this study, we examine in detail the hydrodynamics of American eels (Anguilla rostrata) swimming steadily at 1.4·L·s–1 and compare them with previous results from other fishes. We performed highresolution particle image velocimetry (PIV) to quantify the wake structure, measure the swimming efficiency, and force and power output. The wake consists of jets of fluid that point almost directly laterally, separated by an unstable shear layer that rolls up into two or more vortices over time. Previously, the wake of swimming eels was hypothesized to consist of unlinked vortex rings, resulting from a phase offset between vorticity distributed along the body and vorticity shed at the tail. Our highresolution flow data suggest that the body anterior to the tail tip produces relatively low vorticity, and instead the wake structure results from the instability of the shear layers separating the lateral jets, reflecting pulses of high vorticity shed at the tail tip. We compare the wake structure to large-amplitude elongated body theory and to a previous computational fluid dynamic model and note several discrepancies between the models and the measured values. The wake of steadily swimming eels differs substantially in structure from the wake of previously studied carangiform fishes in that it lacks any significant downstream flow, previously interpreted as signifying thrust. We infer that the lack of downstream flow results from a spatial and temporal balance of momentum removal (drag) and thrust generated along the body, due to the relatively uniform shape of eels. Carangiform swimmers typically have a narrow caudal peduncle, which probably allows them to separate thrust from drag both spatially and temporally. Eels seem to lack this separation, which may explain why they produce a wake with little downstream momentum while carangiform swimmers produce a wake with a clear thrust signature.

The kinematics of swimming frogs have been studied extensively in the past and, based on these results, hypotheses regarding the hydrodynamics of frog swimming can be generated. To test these hypotheses we used digital particle image velocimetry (DPIV) to quantify the flow structure of the wake produced by the feet during the propulsion phase of the kick of surface swimming frogs (Rana pipiens). These frogs use two different gaits, asynchronous and synchronous kicking, and the magnitude of the thrust produced by the feet differs between asynchronous (34±5.4·mN·foot–1) and synchronous kicking (71±13.3·mN·foot–1), as does maximum swimming speed, with higher swimming speed and forces produced during the synchronous kicks. Previous studies have suggested that an interaction between the feet, resulting in a single posteriorly directed fluid jet, as the feet come together at the end of synchronous kicks, may augment force production. Our results show, however, that each foot produces its own distinct vortex ring, in both asynchronous and synchronous kicking of the feet. There is no evidence of a central jet being produced even during powerful synchronous kicks (maximum thrust calculated was 264·mN·foot–1). An alternative mechanism of force production could be the lift-based paddling recently suggested for delta-shaped feet of swimming birds. However, the orientation of the vortex rings generated by the feet is almost perpendicular to the swimming direction for both gaits and there is only a slight asynchrony of the shedding of the distal (start) and proximal (stop) vortex rings, which is different from what would be expected by a dominantly lift-based mechanism. Thus, our results do not support lift as a major mechanism contributing to thrust. Instead, our data support the hypothesis that propulsion is based on drag and acceleration reaction forces where the thrust is generated by separated, but attached, vortex rings on the suction side of the feet, resulting in vortices that are shed behind the frogs during both asynchronous and synchronous kicking.

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.

Swimming and flying animals generate unsteady locomotive forces by delivering net momentum into the fluid wake. Hence, swimming and flying forces can be quantified by measuring the momentum of animal wakes. A recently developed model provides an approach to empirically deduce swimming and flying forces based on the measurement of velocity and vortex added-mass in the animal wake. The model is contingent on the identification of the vortex boundary in the wake. This paper demonstrates the application of that method to a case study quantifying the instantaneous locomotive forces generated by the pectoral fins of the bluegill sunfish (Lepomis macrochirus Rafinesque), measured using digital particle image velocimetry (DPIV). The finite-time Lyapunov exponent (FTLE) field calculated from the DPIV data was used to determine the wake vortex boundary, according to recently developed fluid dynamics theory. Momentum of the vortex wake and its added-mass were determined and the corresponding instantaneous locomotive forces were quantified at discrete time points during the fin stroke. The instantaneous forces estimated in this study agree in magnitude with the time-averaged forces quantified for the pectoral fin of the same species swimming in similar conditions and are consistent with the observed global motion of the animals. A key result of this study is its suggestion that the dynamical effect of the vortex wake on locomotion is to replace the real animal fin with an ‘effective appendage’, whose geometry is dictated by the FTLE field and whose interaction with the surrounding fluid is wholly dictated by inviscid concepts from potential flow theory. Benefits and limitations of this new framework for non-invasive instantaneous force measurement are discussed, and its application to comparative biomechanics and engineering studies is suggested.

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.

In this study, we examine the kinematics and hydrodynamics of the soft dorsal fin in a representative basal teleost, the rainbow trout (Oncorhynchus mykiss), during steady rectilinear locomotion at 0.5–2.0 body lengths (L)·s–1 and during maneuvering. During steady swimming, dorsal fin height and sweep amplitude decrease with increasing speed. The dorsal fin wake, as viewed within a horizontal plane, consists of paired vortices on each side of the body (0.5·L·s–1) or nearly linearly arrayed vortex centers above the body (1.0·L·s–1) with central jet flows directed predominately laterally (lateral:thrust force ratio=5–6). At 2.0·L·s–1, the dorsal fin is no longer recruited to add momentum to the wake. This pattern of decreasing involvement of the trout dorsal fin in thrust production with increasing speed contrasts with the results of our previous study of the soft dorsal fin of sunfish (Lepomis), which is hydrodynamically inactive at low speed and sheds a propulsive vortex wake at higher speed. Yawing maneuvers by trout involve unilateral production of a single vortex ring by the dorsal fin with a strong jet flow oriented almost directly laterally. During steady swimming, interception by the tail of the dorsal fin’s vortical wake and the adipose fin’s non-vortical (drag) wake is hypothesized as a mechanism for enhancing tail thrust. This study provides the first experimental evidence that the plesiomorphic soft dorsal fin of ray- finned fishes acts as an ancillary force generator during axial locomotion. We suggest that the distinction often made between median and paired fin (MPF) propulsion and body and caudal fin (BCF) propulsion in fishes obscures the important role of multiple propulsors acting in a coordinated fashion. Using a combination of anterior median fin oscillation and axial undulation, without continuous paired fin excursions, trout employ an ‘M–BCF’ gait during steady swimming. The primarily lateral orientation of dorsal fin force in trout induces corresponding roll and yaw moments, which must be countered by forces from the caudal, anal and paired fins. Locomotion in trout therefore involves the simultaneous active use of multiple fins, presumably to maintain body stability in the face of environmental perturbations.

Many fish species span two or three orders of magnitude in length during the growth from larvae to adults, and this change may have dramatic consequences for locomotor performance. We measured how the performance of coasting changes over the life history of zebrafish (Danio rerio) and examined the scaling of mechanics underlying this change. Adult zebrafish coast disproportionately further and faster and maintain their speed for a longer duration than do larvae and juveniles. Measurements of drag on tethered dead fish suggest that adult fish operate in an inertial regime by coasting at relatively high Reynolds numbers (Re>1000), and in vivo drag measurements showed adults to operate with a drag coefficient (Cinert 0.024) that was consistent with previously published estimates. However, drag scaled differently at lower Re values than those assumed in previous studies. We found a viscous regime at Re<300, which corresponds to the routine coasting of larvae and juveniles. Despite these changes in hydrodynamics over growth, a mathematical model of coasting mechanics suggests that the disproportionately longer coasting of adults is caused primarily by their large body mass and high speed at the beginning of coasting. We therefore propose that changes in coasting performance with growth are dictated primarily by the scaling of momentum rather than resulting from hydrodynamic changes. These results provide an opportunity for new interpretations of function in the growth and evolution of fish.

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.

Many natural animal movements involve accelerating from a standstill and then stopping. Obstacles in natural environments often limit the straight-line distance available for movement, and decreased braking ability theoretically can limit speed for short distances. Consequently, braking ability can be important for avoiding collisions with obstacles and exploiting resources effectively in complex environments. A presumed morphological correlate of improved braking performance in fish is increased pectoral fin area, because most fish protract these structures as they decelerate. However, the kinematics and modulation of velocity during starting and stopping are poorly understood for most species of fish as well as most species of animals. Thus, for bluegill sunfish Lepomis macrochirus with complete and partially ablated pectoral fins (35% original fin area), we analyzed high speed video recordings (200·images·s–1) of predatory attacks with a start and stop in a short, standardized distance (40·cm). We quantified body displacement, velocity, acceleration, deceleration and several fin angle variables during each feeding. Unexpectedly, several variables including maximum velocity and maximum deceleration (grand means 72·cm·s–1 and –512·cm·s–2, respectively) did not change significantly with reduced pectoral fin area. The average values of braking movements of the median and caudal fins did increase with decreased pectoral fin area but lacked statistically significant differences. The primary mechanism of attaining similar braking performance with decreased area of the pectoral fins was that they were protracted significantly more (mean difference=42°) and with a significantly faster average velocity of protraction. Thus, pectoral fin area appears unlikely to be the primary constraint on braking performance for this particular task.