Convergence of undulatory swimming kinematics across a diversity of fishes.

Fishes exhibit an astounding diversity of locomotor behaviors from classic swimming with their body and fins to jumping, flying, walking, and burrowing. Fishes that use their body and caudal fin (BCF) during undulatory swimming have been traditionally divided into modes based on the length of the propulsive body wave and the ratio of head:tail oscillation amplitude: anguilliform, subcarangiform, carangiform, and thunniform. This classification was first proposed based on key morphological traits, such as body stiffness and elongation, to group fishes based on their expected swimming mechanics. Here, we present a comparative study of 44 diverse species quantifying the kinematics and morphology of BCF-swimming fishes. Our results reveal that most species we studied share similar oscillation amplitude during steady locomotion that can be modeled using a second-degree order polynomial. The length of the propulsive body wave was shorter for species classified as anguilliform and longer for those classified as thunniform, although substantial variability existed both within and among species. Moreover, there was no decrease in head:tail amplitude from the anguilliform to thunniform mode of locomotion as we expected from the traditional classification. While the expected swimming modes correlated with morphological traits, they did not accurately represent the kinematics of BCF locomotion. These results indicate that even fish species differing as substantially in morphology as tuna and eel exhibit statistically similar two-dimensional midline kinematics and point toward unifying locomotor hydrodynamic mechanisms that can serve as the basis for understanding aquatic locomotion and controlling biomimetic aquatic robots.

Swimming behaviour of silver carp (Hypophthalmichthys molitrix) in response to turbulent flow induced by a D-cylinder.

Turbulence is a complex hydraulic phenomenon which commonly occurs in natural streams and fishways. Riverine fish are subjected to heterogeneous flow velocities and turbulence, which may affect their movements and ability to pass the fishways. However, studies focusing on fish response to turbulent flows are lacking for many species. Here we investigate the effects of the turbulence created by a vertical half-cylinder of various diameters (1.9, 2.5, 3.2 and 5.0 cm) on the swimming ability and behaviour of silver carp, Hypophthalmichthys molitrix. The large D-cylinders (3.0 and 5.0 cm) create specific vorticity and reduced velocities areas in their vicinity, which favours flow refuging behaviours (FRBs) and thus increased relative critical swimming speeds (Urcrit , BL/s) of silver carp, by comparison to free-flow conditions and cylinders of smaller diameter (1.9 and 2.5 cm). The flow speed at which silver carp maximized FRBs such as Karman gaiting downstream of the cylinder, holding position in the bow wake or entraining on the side ranged from 40 to 70 cm s-1 , depending on fish body size. When holding station near a cylinder under optimal flow speeds, the distance between the fish and the cylinder is related to the size of the fish, but also to the size of the cylinder and the produced vortices. The optimal holding region in the drag wake of the cylinder ranged from 28 to 40 cm downstream of the centre of the cylinder, depending on the size of the fish. Smaller fish, however, tend to use the reduced velocities areas located in the bow wake of the large cylinders. We hypothesize that fish will display FRBs, including maintaining a Karman gait in turbulent flow, when the ratio of the cylinder diameter to their body length is between 1:3 and 1:4. They also match their tail beat frequency to the vortex shedding frequency of the cylinder. Our results provide a better understanding of how silver carp respond to turbulent flows around physical structures, with implications for the design of nature-like fishways or exclusion devices in both its native and invasive ranges.

Fish-inspired segment models for undulatory steady swimming.

Many aquatic animals swim by undulatory body movements and understanding the diversity of these movements could unlock the potential for designing better underwater robots. Here, we analyzed the steady swimming kinematics of a diverse group of fish species to investigate whether their undulatory movements can be represented using a series of interconnected multi-segment models, and if so, to identify the key factors driving the segment configuration of the models. Our results show that the steady swimming kinematics of fishes can be described successfully using parsimonious models, 83% of which had fewer than five segments. In these models, the anterior segments were significantly longer than the posterior segments, and there was a direct link between segment configuration and swimming kinematics, body shape, and Reynolds number. The models representing eel-like fishes with elongated bodies and fishes swimming at high Reynolds numbers had more segments and less segment length variability along the body than the models representing other fishes. These fishes recruited their anterior bodies to a greater extent, initiating the undulatory wave more anteriorly. Two shape parameters, related to axial and overall body thickness, predicted segment configuration with moderate to high success rate. We found that head morphology was a good predictor of its segment length. While there was a large variation in head segments, the length of tail segments was similar across all models. Given that fishes exhibited variable caudal fin shapes, the consistency of tail segments could be a result of an evolutionary constraint tuned for high propulsive efficiency. The bio-inspired multi-segment models presented in this study highlight the key bending points along the body and can be used to decide on the placement of actuators in fish-inspired robots, to model hydrodynamic forces in theoretical and computational studies, or for predicting muscle activation patterns during swimming.