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Pseudogiant Number Fluctuations and Nematic Order in Microswimmer Suspensions

Ismail El Korde1, Dóra Bárdfalvy1, Jason M. Lewis1, Alexander Morozov2, Cesare Nardini3,4, and Joakim Stenhammar1,*

  • *Contact author: joakim.stenhammar@fkem1.lu.se

Phys. Rev. Lett. 135, 098302 – Published 27 August, 2025

DOI: https://doi.org/10.1103/31q1-hj3n

Abstract

Giant number fluctuations (GNFs), whereby the standard deviation ΔN in the local number of particles ⟨N⟩ grows faster than ⟨N⟩, are a hallmark property of dry active matter systems with orientational order, such as a collection of granular particles on a vibrated plate. This contrasts with momentum-conserving (“wet”) active matter systems, such as suspensions of swimming bacteria, where no theoretical prediction of GNFs exist, although numerous experimental observations of such enhanced fluctuations have been reported. In this Letter, we numerically confirm the emergence of super-Gaussian number fluctuations in a three-dimensional suspension of pusher microswimmers undergoing a transition to collective motion. These fluctuations emerge sharply above the transition, but only for sufficiently large values of the bacterial persistence length ℓp=vs/λ, where vs is the bacterial swimming speed and λ the tumbling rate. Crucially, these “pseudo-GNFs” differ from true GNFs, as they only occur on length scales shorter than the typical size ξ of nematic patches in the collective motion state, which is in turn proportional to the single-swimmer persistence length ℓp. Our results thus suggest that observations of enhanced density fluctuations in biological active matter systems actually represent transient effects that decay away beyond mesoscopic length scales and raises the question to what extent “true” GNFs with universal properties can exist in the presence of fluid flows.

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References (42)

  1. M. C. Marchetti, J. F. Joanny, S. Ramaswamy, T. B. Liverpool, J. Prost, M. Rao, and R. A. Simha, Hydrodynamics of soft active matter, Rev. Mod. Phys. 85, 1143 (2013).
  2. S. Ramaswamy, The mechanics and statistics of active matter, Annu. Rev. Fluid Mech. 1, 323 (2010).
  3. V. Narayan, S. Ramaswamy, and N. Menon, Long-lived giant number fluctuations in a swarming granular nematic, Science 317, 105 (2007).
  4. N. Kumar, H. Soni, S. Ramaswamy, and A. K. Sood, Flocking at a distance in active granular matter, Nat. Commun. 5, 4688 (2014).
  5. J. Deseigne, O. Dauchot, and H. Chaté, Collective motion of vibrated polar disks, Phys. Rev. Lett. 105, 098001 (2010).
  6. H. Chaté, F. Ginelli, and R. Montagne, Simple model for active nematics: Quasi-long-range order and giant fluctuations, Phys. Rev. Lett. 96, 180602 (2006).
  7. H. Chaté, F. Ginelli, G. Grégoire, and F. Raynaud, Collective motion of self-propelled particles interacting without cohesion, Phys. Rev. E 77, 046113 (2008).
  8. S. Ramaswamy, R. A. Simha, and J. Toner, Active nematics on a substrate: Giant number fluctuations and long-time tails, Europhys. Lett. 62, 196 (2003).
  9. J. Toner, Giant number fluctuations in dry active polar fluids: A shocking analogy with lightning rods, J. Chem. Phys. 150, 154120 (2019).
  10. S. Shankar, S. Ramaswamy, and M. C. Marchetti, Low-noise phase of a two-dimensional active nematic system, Phys. Rev. E 97, 012707 (2018).
  11. R. A. Simha and S. Ramaswamy, Hydrodynamic fluctuations and instabilities in ordered suspensions of self-propelled particles, Phys. Rev. Lett. 89, 058101 (2002).
  12. A. Maitra, P. Srivastava, M. C. Marchetti, J. S. Lintuvuori, S. Ramaswamy, and M. Lenz, A nonequilibrium force can stabilize 2D active nematics, Proc. Natl. Acad. Sci. U.S.A. 115, 6934 (2018).
  13. Z. Liu, W. Zeng, X. Ma, and X. Cheng, Density fluctuations and energy spectra of 3D bacterial suspensions, Soft Matter 17, 10806 (2021).
  14. G. Duclos, S. Garcia, H. G. Yevick, and P. Silberzan, Perfect nematic order in confined monolayers of spindle-shaped cells, Soft Matter 10, 2346 (2014).
  15. D. Nishiguchi, K. H. Nagai, H. Chaté, and M. Sano, Long-range nematic order and anomalous fluctuations in suspensions of swimming filamentous bacteria, Phys. Rev. E 95, 020601(R) (2017).
  16. F. Peruani, J. Starruß, V. Jakovljevic, L. Søgaard-Andersen, A. Deutsch, and M. Bär, Collective motion and nonequilibrium cluster formation in colonies of gliding bacteria, Phys. Rev. Lett. 108, 098102 (2012).
  17. H. P. Zhang, A. Be’er, E.-L. Florin, and H. L. Swinney, Collective motion and density fluctuations in bacterial colonies, Proc. Natl. Acad. Sci. U.S.A. 107, 13626 (2010).
  18. D. Saintillan and M. J. Shelley, Emergence of coherent structures and large-scale flows in motile suspensions, J. R. Soc. Interface 9, 571 (2012).
  19. J. Gachelin, A. Rousselet, A. Lindner, and E. Clément, Collective motion in an active suspension of Escherichia coli bacteria, New J. Phys. 16, 025003 (2014).
  20. A. Sokolov and I. S. Aranson, Physical properties of collective motion in suspensions of bacteria, Phys. Rev. Lett. 109, 248109 (2012).
  21. J. Dunkel, S. Heidenreich, K. Dreschner, H. H. Wensink, M. Bär, and R. E. Goldstein, Fluid dynamics of bacterial turbulence, Phys. Rev. Lett. 110, 228102 (2013).
  22. H. H. Wensink, J. Dunkel, S. Heidenreich, K. Drescher, R. E. Goldstein, H. Löwen, and J. M. Yeomans, Meso-scale turbulence in living fluids, Proc. Natl. Acad. Sci. U.S.A. 109, 14308 (2012).
  23. C. Hohenegger and M. J. Shelley, Stability of active suspensions, Phys. Rev. E 81, 046311 (2010).
  24. G. Subramanian and D. L. Koch, Critical bacterial concentration for the onset of collective swimming, J. Fluid Mech. 632, 359 (2009).
  25. J. Stenhammar, C. Nardini, R. W. Nash, D. Marenduzzo, and A. Morozov, Role of correlations in the collective behavior of microswimmer suspensions, Phys. Rev. Lett. 119, 028005 (2017).
  26. D. Bárdfalvy, H. Nordanger, C. Nardini, A. Morozov, and J. Stenhammar, Particle-resolved lattice Boltzmann simulations of 3-dimensional active turbulence, Soft Matter 15, 7747 (2019).
  27. D. Saintillan and M. J. Shelley, Instabilities, pattern formation, and mixing in active suspensions, Phys. Fluids 20, 123304 (2008).
  28. V. Škultéty, C. Nardini, J. Stenhammar, D. Marenduzzo, and A. Morozov, Swimming suppresses correlations in dilute suspensions of pusher microorganisms, Phys. Rev. X 10, 031059 (2020).
  29. V. Škultéty, D. Bárdfalvy, J. Stenhammar, C. Nardini, and A. Morozov, Hydrodynamic instabilities in a two-dimensional sheet of microswimmers embedded in a three-dimensional fluid, J. Fluid Mech. 980, A28 (2024).
  30. D. Bárdfalvy, V. Škultéty, C. Nardini, A. Morozov, and J. Stenhammar, Collective motion in a sheet of microswimmers, Commun. Phys. 7, 93 (2024).
  31. R. W. Nash, R. Adhikari, and M. E. Cates, Singular forces and pointlike colloids in lattice Boltzmann hydrodynamics, Phys. Rev. E 77, 026709 (2008).
  32. R. W. Nash, R. Adhikari, J. Tailleur, and M. E. Cates, Run-and-tumble particles with hydrodynamics: Sedimentation, trapping, and upstream swimming, Phys. Rev. Lett. 104, 258101 (2010).
  33. V. A. Martinez, E. Clément, J. Arlt, C. Douarche, A. Dawson, J. Schwarz-Linek, A. K. Creppy, V. Škultéty, A. N. Morozov, H. Auradou, and W. C. K. Poon, A combined rheometry and imaging study of viscosity reduction in bacterial suspensions, Proc. Natl. Acad. Sci. U.S.A. 117, 2326 (2020).
  34. See Supplemental Material containing finite-size scalings of fluid velocity variance and GNF exponents as well as fits used to extract the correlation lengths in Fig. 3, at http://link.aps.org/supplemental/10.1103/31q1-hj3n for further details.
  35. D. Saintillan and M. J. Shelley, Theory of active suspensions, in Complex Fluids in Biological Systems: Experiment, Theory, and Computation, edited by S. E. Spagnolie (Springer, New York, 2015), pp. 319–355.
  36. B. Martínez-Prat, J. Ignés-Mullol, J. Casademunt, and F. Sagués, Selection mechanism at the onset of active turbulence, Nat. Phys. 15, 362 (2019).
  37. V. Škultéty, Collective motion in microswimmer suspensions, Ph. D. thesis, The University of Edinburgh, 2023.
  38. A. Baskaran and M. C. Marchetti, Statistical mechanics and hydrodynamics of bacterial suspensions, Proc. Natl. Acad. Sci. U.S.A. 106, 15567 (2009).
  39. A. Doostmohammadi, J. Ignés-Mullol, J. Yeomans, and F. Sagués, Active nematics, Nat. Commun. 9, 3246 (2018).
  40. S. Santhosh, M. R. Nejad, A. Doostmohammadi, J. M. Yeomans, and S. P. Thampi, Activity induced nematic order in isotropic liquid crystals, J. Stat. Phys. 180, 699 (2020).
  41. J. de Graaf and J. Stenhammar, Lattice-Boltzmann simulations of microswimmer-tracer interactions, Phys. Rev. E 95, 023302 (2017).
  42. K. Drescher, J. Dunkel, L. H. Cisneros, S. Ganguly, and R. E. Goldstein, Fluid dynamics and noise in bacterial cell-cell and cell-surface scattering, Proc. Natl. Acad. Sci. U.S.A. 108, 10940 (2011).

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