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  • Open Access

How flagellated bacteria wobble

Chen Gui1,*, Mingxin Mao1,2,*,†, Pu Feng3,*, Yuxin Shen3, Xiangjun Gong3,‡, Gerhard Gompper4,§, and Jinglei Hu1,∥

  • *These authors contributed equally to this work.
  • †Contact author: maomx25@mails.tsinghua.edu.cn
  • ‡Contact author: msxjgong@scut.edu.cn
  • §Contact author: g.gompper@fz-juelich.de
  • ∥Contact author: hujinglei@nju.edu.cn

Phys. Rev. Research 8, L012027 – Published 28 January, 2026

DOI: https://doi.org/10.1103/g3w4-5jw8

Abstract

The wobbling motion of flagellated bacteria significantly influences their swimming behavior, yet its dynamics remains poorly understood, primarily due to the difficulty in tracking the 3D orientations of both the flagellar bundle and the cell body. Here, we use hydrodynamics simulations of a mechanical Escherichia coli model in polymer fluids with varying wobbling amplitudes and flagellar anchoring configurations to elucidate this motion. Through Euler angle analysis, we resolve the three components of wobbling: precession, nutation, and spin. Our findings challenge the common assumption that peritrichous bacteria undergo complete cycles of body spin. Notably, we uncover a small-amplitude nutation with a well-defined period on the order of a few milliseconds. We also identify two distinct precession modes. To interpret these findings, we develop a theoretical framework based on Lagrangian mechanics.

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Physics Subject Headings (PhySH)

Corrections

10 March, 2026

Correction: The equal contribution footnote for the first three authors was missing at publication and has been inserted.

Article Text

Supplemental Material

References (49)

  1. H. C. Berg, E. Coli in Motion (Springer, New York, 2004).
  2. N. C. Darnton, L. Turner, S. Rojevsky, and H. C. Berg, On torque and tumbling in swimming Escherichia coli, J. Bacteriol. 189, 1756 (2007).
  3. N. Watari and R. G. Larson, Swimming hydrodynamics of a run-and-tumble bacterium with helical flagella, Biophys. J. 98, 160A (2010).
  4. V. A. Martinez, J. Schwarz-Linek, M. Reufer, L. G. Wilson, A. N. Morozov, and W. C. K. Poon, Flagellated bacterial motility in polymer solutions, Proc. Natl. Acad. Sci. USA 111, 17771 (2014).
  5. B. Liu, M. Gulino, M. Morse, J. X. Tang, T. R. Powers, and K. S. Breuer, Helical motion of the cell body enhances Caulobacter crescentus motility, Proc. Natl. Acad. Sci. USA 111, 11252 (2014).
  6. J. Hu, M. Yang, G. Gompper, and R. G. Winkler, Modelling the mechanics and hydrodynamics of swimming E. coli, Soft Matter 11, 7867 (2015).
  7. A. E. Patteson, A. Gopinath, M. Goulian, and P. E. Arratia, Running and tumbling with E. coli in polymeric solutions, Sci. Rep. 5, 15761 (2015).
  8. A. Zöttl and J. M. Yeomans, Enhanced bacterial swimming speeds in macromolecular polymer solutions, Nat. Phys. 15, 554 (2019).
  9. S. Kamdar, S. Shin, P. Leishangthem, L. F. Francis, X. Xu, and X. Cheng, The colloidal nature of complex fluids enhances bacterial motility, Nature (London) 603, 819 (2022).
  10. W. R. DiLuzio, L. Turner, M. Mayer, P. Garstecki, D. B. Weibel, H. C. Berg, and G. M. Whitesides, Escherichia coli swim on the right-hand side, Nature (London) 435, 1271 (2005).
  11. E. Lauga, W. R. DiLuzio, G. M. Whitesides, and H. A. Stone, Swimming in circles: Motion of bacteria near solid boundaries, Biophys. J. 90, 400 (2006).
  12. A. P. Berke, L. Turner, H. C. Berg, and E. Lauga, Hydrodynamic attraction of swimming microorganisms by surfaces, Phys. Rev. Lett. 101, 038102 (2008).
  13. R. Di Leonardo, D. Dell’Arciprete, L. Angelani, and V. Iebba, Swimming with an image, Phys. Rev. Lett. 106, 038101 (2011).
  14. M. Molaei, M. Barry, R. Stocker, and J. Sheng, Failed escape: Solid surfaces prevent tumbling of Escherichia coli, Phys. Rev. Lett. 113, 068103 (2014).
  15. J. Hu, A. Wysocki, R. G. Winkler, and G. Gompper, Physical sensing of surface properties by microswimmers—Directing bacterial motion via wall slip, Sci. Rep. 5, 9586 (2015).
  16. S. Bianchi, F. Saglimbeni, and R. Di Leonardo, Holographic imaging reveals the mechanism of wall entrapment in swimming bacteria, Phys. Rev. X 7, 011010 (2017).
  17. D. Cao, M. Dvoriashyna, S. Liu, E. Lauga, and Y. Wu, Reduced surface accumulation of swimming bacteria in viscoelastic polymer fluids, Proc. Natl. Acad. Sci. USA 119, e2212078119 (2022).
  18. X. Chen, X. Dong, A. Be'er, H. L. Swinney, and H. P. Zhang, Scale-invariant correlations in dynamic bacterial clusters, Phys. Rev. Lett. 108, 148101 (2012).
  19. C. Chen, S. Liu, X.-Q. Shi, H. Chaté, and Y. Wu, Weak synchronization and large-scale collective oscillation in dense bacterial suspensions, Nature (London) 542, 210 (2017).
  20. S. Liu, S. Shankar, M. C. Marchetti, and Y. Wu, Viscoelastic control of spatiotemporal order in bacterial active matter, Nature (London) 590, 80 (2021).
  21. Y. Hyon, Marcos, T. R. Powers, R. Stocker, and H. C. Fu, The wiggling trajectories of bacteria, J. Fluid Mech. 705, 58 (2012).
  22. M. A. Constantino, M. Jabbarzadeh, H. C. Fu, and R. Bansil, Helical and rod-shaped bacteria swim in helical trajectories with little additional propulsion from helical shape, Sci. Adv. 2, e1601661 (2016).
  23. S. Bianchi, F. Saglimbeni, A. Lepore, and R. Di Leonardo, Polar features in the flagellar propulsion of E. coli bacteria, Phys. Rev. E 91, 062705 (2015).
  24. H. Zheng, N. Yan, W. Feng, Y. Liu, H. Luo, and G. Jing, Swimming of buoyant bacteria in quiescent medium and shear flows, Langmuir 39, 4224 (2023).
  25. T.-W. Su, L. Xue, and A. Ozcan, High-throughput lensfree 3D tracking of human sperms reveals rare statistics of helical trajectories, Proc. Natl. Acad. Sci. USA 109, 16018 (2012).
  26. M. Qi, X. Gong, B. Wu, and G. Zhang, Landing dynamics of swimming bacteria on a polymeric surface: Effect of surface properties, Langmuir 33, 3525 (2017).
  27. S. M. Mousavi, G. Gompper, and R. G. Winkler, Wall entrapment of peritrichous bacteria: A mesoscale hydrodynamics simulation study, Soft Matter 16, 4866 (2020).
  28. The addition of polymer chains allows for systematic tuning of the angle θ, while leaving flagellar anchoring and bundling largely unaffected, as illustrated in Fig.  3. The increased fluid viscosity upon polymer addition also facilitates the visual inspection of precession and nutation (Fig.  2). Although one could simulate a population of bacteria with varying flagellar number and positioning to achieve different wobbling amplitudes, this approach is less efficient than adding polymer chains.
  29. See Supplemental Material at http://link.aps.org/supplemental/10.1103/g3w4-5jw8 for details, which includes Refs.  [26, 30, 31, 32, 33, 34, 35, 36, 37, 38, 39, 40, 41, 42, 43, 44, 45].
  30. A. Malevanets and R. Kapral, Mesoscopic model for solvent dynamics, J. Chem. Phys. 110, 8605 (1999).
  31. G. Gompper, T. Ihle, D. M. Kroll, and R. G. Winkler, Multi-particle collision dynamics: A particle-based mesoscale simulation approach to the hydrodynamics of complex fluids, Adv. Polym. Sci. 221, 1 (2009).
  32. H. Noguchi and G. Gompper, Shape transitions of fluid vesicles and red blood cells in capillary flows, Proc. Natl. Acad. Sci. USA 102, 14159 (2005).
  33. S. Sahoo, S. P. Singh, and S. Thakur, Role of viscoelasticity on the dynamics and aggregation of chemically active sphere-dimers, Phys. Fluids 33, 017120 (2021).
  34. M. Chen, Z. Lin, M. Xuan, X. Lin, M. Yang, L. Dai, and Q. He, Programmable dynamic shapes with a swarm of light-powered colloidal motors, Angew. Chem. Intl. Ed. 60, 16674 (2021).
  35. T. Li, Z. Liu, J. Hu, L. Chen, T. Chen, Q. Tang, B. Yu, B. Zhao, C. Mao, and M. Wan, A universal chemotactic targeted delivery strategy for inflammatory diseases, Adv. Mater. 34, 2206654 (2022).
  36. H. Yamakawa, Helical Wormlike Chains in Polymer Solutions (Springer, Berlin, 1997).
  37. R. Vogel and H. Stark, Force-extension curves of bacterial flagella, Eur. Phys. J. E 33, 259 (2010).
  38. P. J. A. Janssen and M. D. Graham, Coexistence of tight and loose bundled states in a model of bacterial flagellar dynamics, Phys. Rev. E 84, 011910 (2011).
  39. S. Y. Reigh, R. G. Winkler, and G. Gompper, Synchronization and bundling of anchored bacterial flagella, Soft Matter 8, 4363 (2012).
  40. R. M. Berry and H. C. Berg, Absence of a barrier to backwards rotation of the bacterial flagellar motor demonstrated with optical tweezers, Proc. Natl. Acad. Sci. USA 94, 14433 (1997).
  41. M. Qi, Q. Song, J. Zhao, C. Ma, G. Zhang, and X. Gong, Three-dimensional bacterial behavior near dynamic surfaces formed by degradable polymers, Langmuir 33, 13098 (2017).
  42. F. C. Cheong, B. J. Krishnatreya, and D. G. Grier, Strategies for three-dimensional particle tracking with holographic video microscopy, Opt. Express 18, 13563 (2010).
  43. H. Goldstein, C. P. Poole, and J. L. Safko, Classical Mechanics (Pearson, Upper Saddle River, NJ, 2001), 3rd ed.
  44. C. Yang, C. Chen, Q. Ma, L. Wu, and T. Song, Dynamic model and motion mechanism of magnetotactic bacteria with two lateral flagellar bundles, J. Bionic Eng. 9, 200 (2012).
  45. J. Lighthill, Flagellar hydrodynamics, SIAM Rev. 18, 161 (1976).
  46. The overall rotation of the body is primarily due to precession, while the overall rotation of the flagellar bundle is dominated by spin. These two overall rotations are in opposite directions.
  47. The Python script and exemplary trajectory file for Euler angle analysis are available at https://doi.org/10.6084/m9.figshare.30121015.
  48. There are no such physical constraints in uniflagellated bacteria. The spin of body at a rate of about 30 Hz has been reported for Caulobacter crescentus, when the single flagellum pushes the cell forward [5].
  49. Our mesoscale hydrodynamics simulations that automatically include thermal fluctuations reveal that the time averages of the two instantaneous quantities, L¯z and L¯3, are constants.

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