The importance of body stiffness in undulatory propulsion1

Abstract

SYNOPSIS. During steady swimming in fish, the dynamic form taken by the axial undulatory wave may depend on the bending stiffness of the body. Previous studies have suggested the hypothesis that fish use their muscles to modulate body stiffness. In order to expand the theoretical and experimental tools available for testing this hypothesis, we explored the relationship between body stiffness, muscle activity, and undulatory waveform in the mechanical context of dynamically bending beams. We propose that fish minimize the mechanical cost of bending by increasing their body stiffness, which would allow them to tune their body's natural frequency to match the tailbeat frequency at a given swimming speed. A review of the literature reveals that the form of the undulatory wave, as measured by propulsive wavelength, is highly variable within species, a result which calls into question the use of propulsive wavelength as a species-specific indicator of swimming mode. At the same time, the smallest wavelength within a species is inversely proportional to the number of vertebrae across taxa (r2 = 0.21). In order to determine if intact fish bodies are capable of increasing bending stiffness, we introduce a method for stimulating muscle in the body of a dead fish while it is being cyclically bent at physiological frequencies. The bending moment (N m) and angular displacement (radians) are measured during dynamic bending with and without muscle stimulation. Initial results from these wholebody work loops demonstrate that largemouth bass possess the capability to increase body stiffness by using their muscles to generate negative mechanical work.