A B S T R A C TMany fish swim using body undulations to generate thrust and maneuver in three dimensions. The pattern of body bending during steady rectilinear locomotion has similar general characteristics in many fishes and involves a wave of increasing amplitude passing from the head region toward the tail. While great progress has been made in understanding the mechanics of undulatory propulsion in fishes, the inability to control and precisely alter individual parameters such as oscillation frequency, body shape, and body stiffness, and the difficulty of measuring forces on freely swimming fishes have greatly hampered our ability to understand the fundamental mechanics of the undulatory mode of locomotion in aquatic systems. In this paper, we present the use of a robotic flapping foil apparatus that allows these parameters to be individually altered and forces measured on self-propelling flapping flexible foils that produce a wave-like motion very similar to that of freely swimming fishes. We use this robotic device to explore the effects of changing swimming speed, foil length, and foil-trailing edge shape on locomotor hydrodynamics, the cost of transport, and the shape of the undulating foil during locomotion. We also examine the passive swimming capabilities of a freshly dead fish body. Finally, we model fin-fin interactions in fishes using dual-flapping foils and show that thrust can be enhanced under correct conditions of foil phasing and spacing as a result of the downstream foil making use of vortical energy released by the upstream foil.

SUMMARYThe American lobster (Homarus americanus) displays a diverse set of locomotory behaviours that includes tail flips, walking and paddling. Paddling is carried out by the four pairs of paddle-shaped pleopods on the ventral abdomen. Although it is recognized that pleopod-generated fluid flows have some locomotory role in adults, reports on their relative importance in locomotion are inconsistent. This paper integrates experimental kinematics and hydrodynamics of lobster pleopod beating to determine the mechanism and magnitude of pleopod force production. A kinematic analysis of pleopod beating in live lobsters showed that the pleopods execute an adlocomotory metachronal beating pattern. We modelled in vivo pleopod kinematics with a set of simple trigonometric functions, and used these functions to program a mechanical lobster model consisting of motor-driven pleopods on a lobster abdomen exoskeleton. Based on flow visualizations obtained from applying particle image velocimetry to the lobster model, we propose that the unsteady metachronal kinematics of the pleopods can maximize thrust by exploiting forces arising from individual pleopod activity and interactions among adjacent pairs. The pleopods continuously entrain fluid surrounding the lobster and create a caudally directed fluid jet oriented parallel to the substratum. Inputting wake morphology and velocity data into a simplified model for steady jet thrust showed that the pleopods of the lobster model produced 27-54 mN of thrust, which is comparable to the propulsive forces generated by other proficient swimmers. These results suggest that lobster pleopods are capable of producing forces of a magnitude that could assist the walking legs in forward propulsion.

Anguilliform mode swimmers pass waves of lateral bending down their elongate bodies to propel forward. Hagfishes (Myxinidae) are classified as anguilliform swimmers, but their unique habits and reduced morphology—including a flexible body lacking a vertebral column—have the potential to translate into unique swimming behaviour within this broad classification. Their roles as active scavengers and hunters can require considerable bouts of swimming, yet quantitative data on hagfish locomotion are limited. Here, we aim to provide a more complete mechanistic understanding of hagfish swimming by quantifying whole-body kinematics of steady swimming in Pacific hagfish (Eptatretus stoutii (Lockington, 1878)) and Atlantic hagfish (Myxine glutinosa L., 1758), species from the two main lineages of Myxinidae. We analyzed high-speed video of hagfishes swimming at voluntary swim speeds and found that both species swim using high-amplitude undulatory waves. Swim speed is generally frequency-modulated, but patterns in wave speed, wavelength, and amplitude along the body and across swim speeds are variable, implying versatile mechanisms for the control of swim speed in these highly flexible fishes. We propose mechanistic explanations for this kinematic variability and compare hagfish with other elongate swimmers, demonstrating that the hagfish’s rich locomotory repertoire adds variety to the already diverse set of locomotory kinematics found in anguilliform swimmers.

Physical models enable researchers to systematically examine complex and dynamic mechanisms of underwater locomotion in ways that would be challenging with freely swimming animals. Previous research on undulatory locomotion, for example, has used rectangular flexible panels that are effectively two-dimensional as proxies for the propulsive surfaces of swimming fishes, but these bear little resemblance to the bodies of elongate eel-like swimming animals. In this paper we use a polyurethane rod (round cross-section) and bar (square cross-section) to represent the body of a swimming Pacific hagfish (Eptatretus stoutii). We actuated the rod and bar in both heave and pitch using a mechanical controller to generate a propulsive wave at frequencies between 0.5 and 2.5 Hz. We present data on (1) how kinematic swimming patterns change with driving frequency in these elongate fish-like models, (2) the thrust-generating capability of these simple models, (3) how forces and work done during propulsion compare between cross-sectional shapes, (4) the wake flow patterns in these swimming models using particle image velocimetry. We also contrast kinematic and hydrodynamic patterns produced by bar and rod models to comparable new experimental data on kinematics and wake flow patterns from freely swimming hagfish. Increasing the driving frequency of bar and rod models reduced trailing edge amplitude and wavelength, and above 2 Hz a nodal point appeared in the kinematic wave. Above 1 Hz, both the rod and bar generated net thrust, with the work per cycle reaching a minimum at 1.5 Hz, and the bar always requiring more work per cycle than the rod. Wake flow patterns generated by the swimming rod and bar included clearly visible lateral jets, but not the caudolaterally directed flows seen in the wakes from freely swimming hagfish.

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