Beyond fish in formation: A two-tier approach for biomechanical studies of collective movement

Despite much of the literature perceiving fish schooling as an organized system with a focus on fixed formations for theoretical analyses, experimental observations suggest that frequent positional rearrangement commonly occurs. Previous studies have also demonstrated that fish schools reduce locomotor costs relative to individuals swimming alone. This introduces an intriguing dichotomy. How can individual fish within schools exhibit dynamic interactions while also saving energy? We hypothesize that schooling dynamics are the result of positional and kinematic modulation of individuals responding to fluid dynamic stimuli from the movement of neighbouring individuals. We propose a two-tier approach to studying kinematic modulation within fish schools. First, quantification of the variation of individual movement in a school relative to that of a solitary individual uses an analytical pipeline combining artificial-intelligence-enabled tracking and video processing. Second, the study of kinematic modulation in response to hydrodynamic stimuli uses a mechanical flapping mechanism coupled with an enclosure to control fish position. We discovered that fish in schools exhibit higher levels of positional and kinematic modulation than individuals swimming alone. Fish swimming in enclosures can robustly respond to fluid stimuli from either a simple robotic fish or other fish located in proximity. This two-tier approach allows high-resolution analysis of positional and kinematic modulation within fish schools and their impacts on energy conservation resulting from collective movement.