Undulatory fish swimming: from muscles to flow

Abstract

AbstractUndulatory swimming is employed by many fish for routine swimming and extended sprints. In this biomechanical review, we address two questions: (i) how the fish's axial muscles power swimming; and (ii) how the fish's body and fins generate thrust. Fish have adapted the morphology of their axial musculature for high power output and efficiency. All but the superficial muscle fibres are arranged along curved trajectories, and the myomeres form nested cones. Two conflicting performance goals shape the fibre trajectories of the axial muscles. Maximum power output requires that all fibres contract uniformly. In a bending fish, uniform contraction in a single myomere can be ensured by curved fibre trajectories. However, uniform strain is only desirable if all muscle fibres have the same contractile properties. The fish needs several muscle‐fibre types that generate maximum power at different contraction speeds to ensure effective muscle power generation across a range of swimming speeds. Consequently, these different muscle‐fibre types are better served by non‐uniform contractions. High power output at a range of swimming speeds requires that muscle fibres with the same contractile properties contract uniformly. The ensuing helical fibre trajectories require cone‐shaped myomeres to reduce wasteful internal deformation of the entire muscle when it contracts. It can be shown that the cone‐shaped myomeres of fish can be explained by two design criteria: uniform contraction (uniform strain hypothesis) and minimal internal deformation (mechanical stability hypothesis). So far, only the latter hypothesis has found strong support. The contracting muscle causes the fish body to undulate. These body undulations interact with the surrounding water to generate thrust. The resulting flow behind the swimming fish forms vortex rings, whose arrangement reflects the fish's swimming performance. Anguilliform swimmers shed individual vortex rings during steady swimming. Carangiform swimmers shed a connected chain of vortex rings. The currently available sections through the total flow fields are often not an honest representation of the total momentum in the water – the wake of carangiform swimmers shows a net backward momentum without the fish accelerating – suggesting that our current picture of the generated flow is incomplete. To accelerate, undulatory swimmers decrease the angle of the vortex rings with the mean path of motion, which is consistent with an increased rate of backward momentum transfer. Carangiform swimmers also enlarge their vortex rings to accelerate and to swim at a higher speed, while eel, which are anguilliform swimmers, shed stronger vortex rings.