Wildebeest herds on rolling hills: Flocking on arbitrary curved surfaces
Abstract
The collective behavior of active agents, whether herds of wildebeest or microscopic actin filaments propelled by molecular motors, is an exciting frontier in biological and soft matter physics. Almost three decades ago, Toner and Tu developed a continuum theory of the collective action of flocks, or herds, that helped launch the modern field of active matter. One challenge faced when applying continuum active matter theories to living phenomena is the complex geometric structure of biological environments. Both macroscopic and microscopic herds move on asymmetric curved surfaces, like undulating grass plains or the surface layers of cells or embryos, which can render problems analytically intractable. In this paper, we present a formulation of the Toner-Tu flocking theory that uses the finite element method to solve the governing equations on arbitrary curved surfaces. First, we test the developed formalism and its numerical implementation in channel flow with scattering obstacles and on cylindrical and spherical surfaces, comparing our results to analytical solutions. We then progress to surfaces with arbitrary curvature, moving beyond previously accessible problems to explore herding behavior on a variety of landscapes. This approach allows the investigation of transients and dynamic solutions not revealed by analytic methods. It also enables versatile incorporation of new geometries and boundary conditions and efficient sweeps of parameter space. Looking forward, the paper presented here lays the groundwork for a dialogue between Toner-Tu theory and data on collective motion in biologically relevant geometries, from drone footage of migrating animal herds to movies of microscopic cytoskeletal flows within cells.
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Published by the American Physical Society under the terms of the Creative Commons Attribution 4.0 International license. Further distribution of this work must maintain attribution to the author(s) and the published article's title, journal citation, and DOI.
Acknowledgement
We are deeply grateful to a number of generous colleagues who have carefully described their thinking and work on the formulation of continuum descriptions on surfaces, and to others who have provided invaluable feedback on this paper. We especially thank A. Agarawal, M. Arroyo, D. Bensimon, M. Bowick, M. Deserno, S. Hirokawa, G. Huber, A. Kitaev (who showed us how to solve the half-space problem), J. Kondev, E. Korkmazhan, D. Krishnamurthy, K. Mandadapu, M. Mani, C. Marchetti, W. Melton, A. Mietke, P. Nelson, S. Nissen, M. Prakash, S. Ramaswamy, S. Shankar, M. Shelley, S. Takatori, J. Toner, Y. Tu, and V. Vitelli. We thank N. Orme for his work on Figs. 1–3. This paper was supported by a Damon Runyon Fellowship Award (C.L.H.), a Burroughs Wellcome Career Award at the Scientific Interface (C.L.H.), NIH R35GM130332 (A.R.D.), an HHMI Faculty Scholar Award (A.R.D.), NIH MIRA 1R35GM118043 (R.P.), and the Chan Zuckerberg Biohub (R.P.). C.L.H. is a Damon Runyon Fellow supported by the Damon Runyon Cancer Research Foundation (DRG-2375-19).
Conflict of Interest
The authors declare that they have no competing interests.
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Additional details
- ISSN
- 2470-0053
- Damon Runyon Cancer Research Foundation
- DRG-2375-19
- Burroughs Wellcome Fund
- National Institutes of Health
- R35GM130332
- Howard Hughes Medical Institute
- National Institutes of Health
- 1R35GM118043
- Chan Zuckerberg Initiative (United States)
- Caltech groups
- Division of Biology and Biological Engineering