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For over a century, scientists have sought to understand how fish orient against an incoming flow, even without visual and flow cues. Here, we elucidate a potential hydrodynamic mechanism of rheotaxis through the study of the bidirectional coupling between fish and the surrounding fluid. By modeling a fish as a vortex dipole in an infinite channel with an imposed background flow, we establish a planar dynamical system for the cross-stream coordinate and orientation. The system dynamics captures the existence of a critical flow speed for fish to successfully orient while performing cross-stream, periodic sweeping movements. Model predictions are examined in the context of experimental observations in the literature on the rheotactic behavior of fish deprived of visual and lateral line cues. The crucial role of bidirectional hydrodynamic interactions unveiled by this model points at an overlooked limitation of existing experimental paradigms to study rheotaxis in the laboratory. 

Mathematical models promise new insights into the mechanisms underlying the emergence of collective behaviour in fish. Here, we establish a mathematical model to examine collective behaviour of zebrafish, a popular animal species in preclinical research. The model accounts for social and hydrodynamic interactions between individuals, along with the burst-and-coast swimming style of zebrafish. Each fish is described as a system of coupled stochastic differential equations, which govern the time evolution of their speed and turn rate. Model parameters are calibrated using experimental observations of zebrafish pairs swimming in a shallow water tank. The model successfully captures the main features of the collective response of the animals, by predicting their preference to swim in-line, with one fish leading and the other trailing. During in-line swimming, the animals share the same orientation and keep a distance from each other, owing to hydrodynamic repulsion. Hydrodynamic interaction is also responsible for an increase in the speed of the pair swimming in-line. By linearizing the equations of motion, we demonstrate local stability of in-line swimming to small perturbations for a wide range of model parameters. Mathematically backed results presented herein support the application of dynamical systems theory to unveil the inner workings of fish collective behaviour.