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Fish often swim in crystallized group formations (schooling) and orient themselves against the incoming flow (rheotaxis). At the intersection of these two phenomena, we investigate the emergence of unique schooling patterns through passive hydrodynamic mechanisms in a fish pair, the simplest subsystem of a school. First, we develop a fluid dynamics-based mathematical model for the positions and orientations of two fish swimming against a flow in an infinite channel, modelling the effect of the self-propelling motion of each fish as a point-dipole. The resulting system of equations is studied to gain an understanding of the properties of the dynamical system, its equilibria and their stability. The system is found to have five types of equilibria, out of which only upstream swimming in in-line and staggered formations can be stable. A stable near-wall configuration is observed only in limiting cases. It is shown that the stability of these equilibria depends on the flow curvature and streamwise interfish distance, below critical values of which, the system may not have a stable equilibrium. The study reveals that simply through passive fluid dynamics, in the absence of any other feedback/sensing, we can justify rheotaxis and the existence of stable in-line and staggered schooling configurations.
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Collective phase transitions in confined fish schools
The collective patterns that emerge in schooling fish are often analyzed using models of self-propelled particles in unbounded domains. However, while schooling fish in both field and laboratory settings interact with domain boundaries, these effects are typically ignored. Here, we propose a model that incorporates geometric confinement, by accounting for both flow and wall interactions, into existing data-driven behavioral rules. We show that new collective phases emerge where the school of fish “follows the tank wall” or “double mills.” Importantly, confinement induces repeated switching between two collective states, schooling and milling. We describe the group dynamics probabilistically, uncovering bistable collective states along with unintuitive bifurcations driving phase transitions. Our findings support the hypothesis that collective transitions in fish schools could occur spontaneously, with no adjustment at the individual level, and opens venues to control and engineer emergent collective patterns in biological and synthetic systems that operate far from equilibrium.
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- Award ID(s):
- 2100209
- PAR ID:
- 10609564
- Publisher / Repository:
- PNAS
- Date Published:
- Journal Name:
- Proceedings of the National Academy of Sciences
- Volume:
- 121
- Issue:
- 44
- ISSN:
- 0027-8424
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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- Free Publicly Accessible Full Text
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Accepted Manuscript1.0
- Journal Article:
- https://doi.org/10.1073/pnas.2406293121
