Go reconfigure: how fish change shape as they swim and evolve

The bodies of fish change shape over propulsive, behavioral, developmental, and evolutionary time scales, a general phenomenon that we call "reconfiguration". Undulatory, postural, and form-reconfiguration can be distinguished, studied independently, and examined in terms of mechanical interactions and evolutionary importance. Using a combination of live, swimming fishes and digital robotic fish that are autonomous and self-propelled, we examined the functional relation between undulatory and postural reconfiguration in forward swimming, backward swimming, and yaw turning. To probe how postural and form reconfiguration interact, the yaw turning of leopard sharks was examined using morphometric and kinematic analyses. To test how undulatory reconfiguration might evolve, the digital robotic fish were subjected to selection for enhanced performance in a simulated ecology in which each individual had to detect and move towards a food source. In addition to the general issue of reconfiguration, these investigations are united by the fact that the dynamics of undulatory and postural reconfigurations are predicted to be determined, in part, by the structural stiffness of the fish's body. Our method defines undulatory reconfiguration as the combined, point-by-point periodic motion of the body, leaving postural reconfiguration as the combined deviations from undulatory reconfiguration. While undulatory reconfiguration appears to be the sole or primary propulsive driver, postural reconfiguration may contribute to propulsion in hagfish and it is correlated with differences in forward, and backward, swimming in lamprey. Form reconfigures over developmental time in leopard sharks in a manner that is consistent with an allometric scaling theory in which structural stiffness of the body is held constant. However, correlation of a form proxy for structural stiffness of the body suggests that body stiffness may scale in order to limit maximum postural reconfiguration during routine yaw turns. When structural stiffness and undulatory frequency are modeled as determining the tail's undulatory wave speed, both factors evolve under selection for enhanced foraging behavior in the digital fish-like robots. The methods used in making these distinctions between kinds of reconfiguration have broad applicability in fish biology, especially for quantifying complex motor behaviors in the wild and for simulating selection on behavior that leads to directional evolution of functional phenotypes.