pith. sign in

arxiv: 2607.01547 · v1 · pith:7VYYLKPCnew · submitted 2026-07-01 · ❄️ cond-mat.soft · physics.flu-dyn

Mixing induced by microswimmers as probed by mutual information

Pith reviewed 2026-07-03 17:57 UTC · model grok-4.3

classification ❄️ cond-mat.soft physics.flu-dyn
keywords microswimmersmixing efficiencymutual informationsquirmer modeladvection-diffusionhydrodynamic interactionsactive matterfluid mixing
0
0 comments X

The pith

Mutual information quantifies mixing by microswimmers and shows an optimal finite squirmer parameter balancing translation and flow.

A machine-rendered reading of the paper's core claim, the machinery that carries it, and where it could break.

The paper establishes that mutual information works as a global measure of mixing efficiency in the complex flows created by microswimmers. Numerical solution of the squirmer flow fields and the advection-diffusion equation for tracers shows that aggregation of swimmers suppresses mixing while positional and orientational disorder enhances it. At fixed energy dissipation, efficiency depends non-monotonically on the squirmer parameter and reaches a maximum at a finite value set by the competition between swimmer translation and dipolar flow generation. Hydrodynamic interactions favor pushers over pullers. Mutual information itself decays in three stages: an early diffusion-dominated regime, an intermediate advection-enhanced regime, and a late relaxation stage limited by system size.

Core claim

Mutual information, previously tested only in simplified models, applies directly to complex microswimmer flows and demonstrates that mixing efficiency is subject to a trade-off between generating strong shear and achieving good dispersion across the domain. In the two-dimensional confined squirmer model this trade-off produces an optimal finite squirmer parameter and favors disordered configurations over aggregates.

What carries the argument

Mutual information computed from the evolving spatial distribution of tracers that obey the advection-diffusion equation in the velocity field of two-dimensional squirmers.

If this is right

  • Swimmer aggregation suppresses mixing while positional and orientational disorder enhances it.
  • At fixed energy dissipation, mixing efficiency reaches a maximum at a finite squirmer parameter.
  • With hydrodynamic interactions included, pushers mix more efficiently than pullers.
  • The decay of mutual information occurs in three distinct stages governed successively by diffusion, advection, and system size.

Where Pith is reading between the lines

These are editorial extensions of the paper, not claims the author makes directly.

  • The same mutual-information diagnostic could be applied to three-dimensional swimmer suspensions to test whether the shear-dispersion trade-off persists.
  • The results suggest design rules for microfluidic mixers that use controlled distributions of artificial microswimmers.
  • Biological microswimmers may have evolved parameters near the reported optimum to improve nutrient dispersal or collective transport.
  • The three-stage decay pattern may appear in other confined active-matter systems and could be checked with minimal models lacking hydrodynamics.

Load-bearing premise

The two-dimensional confined squirmer model together with numerical solution of the advection-diffusion equation captures the essential physics of mixing by real microswimmers.

What would settle it

A three-dimensional simulation or laboratory experiment with real microswimmers in which mutual information fails to show the reported non-monotonic dependence on squirmer parameter or the three-stage temporal decay would falsify the claim that the measure works equally well in complex flows.

Figures

Figures reproduced from arXiv: 2607.01547 by Andrej Vilfan, Ramin Golestanian, Yihong Shi, Yuto Hosaka.

Figure 1
Figure 1. Figure 1: FIG. 1. Representative streamlines of the regularized confined velocity field generated by a single squirmer placed at the center [PITH_FULL_IMAGE:figures/full_fig_p004_1.png] view at source ↗
Figure 2
Figure 2. Figure 2: FIG. 2. Time evolution of the probability density [PITH_FULL_IMAGE:figures/full_fig_p005_2.png] view at source ↗
Figure 3
Figure 3. Figure 3: FIG. 3. Velocity fields and mixing efficiency for systems of four non-interacting stresslets in a square domain of side length [PITH_FULL_IMAGE:figures/full_fig_p006_3.png] view at source ↗
Figure 4
Figure 4. Figure 4: FIG. 4. Mixing efficiency for different swimmer types characterized by the squirmer parameter [PITH_FULL_IMAGE:figures/full_fig_p007_4.png] view at source ↗
Figure 5
Figure 5. Figure 5: FIG. 5. Mixing in systems with different square sizes at the fixed number density of non-interacting swimmers. (a) Example [PITH_FULL_IMAGE:figures/full_fig_p008_5.png] view at source ↗
Figure 6
Figure 6. Figure 6: FIG. 6. Velocity fields of fundamental singularities in two [PITH_FULL_IMAGE:figures/full_fig_p010_6.png] view at source ↗
Figure 7
Figure 7. Figure 7: FIG. 7. Image construction for solving the Stokeslet in a [PITH_FULL_IMAGE:figures/full_fig_p011_7.png] view at source ↗
Figure 8
Figure 8. Figure 8: FIG. 8. Reflection rule for a swimmer reaching the wall. The [PITH_FULL_IMAGE:figures/full_fig_p013_8.png] view at source ↗
read the original abstract

We investigate fluid mixing induced by microswimmers using mutual information as a global, information-theoretic measure of mixing efficiency. For a two-dimensional squirmer model in a confined domain, we compute numerically the swimmer-generated flows and solve the advection-diffusion equation for the transport of tracer particles in the fluid. We show that the spatial distribution of swimmers strongly affects mixing, which is suppressed by swimmer aggregation and enhanced by positional and orientational disorder. At fixed energy dissipation, mixing efficiency depends non-monotonically on the squirmer parameter, with an optimal finite value arising from the balance between swimmer translation and dipolar flow generation. When hydrodynamic interactions are included, pushers outperform pullers. The mutual information as a function of time decays in three stages: an initial diffusion-dominated stage, an intermediate advection enhanced regime, and a final relaxation stage controlled by system size. Our results demonstrate that mutual information, previously validated as a measure of mixing efficiency only in simplified model systems, can equally be used in complex flows. Its application reveals that mixing by microswimmers is subject to a trade-off between the generation of strong shear flows and achieving optimal dispersion across the fluid domain.

Editorial analysis

A structured set of objections, weighed in public.

Desk editor's note, referee report, simulated authors' rebuttal, and a circularity audit. Tearing a paper down is the easy half of reading it; the pith above is the substance, this is the friction.

Referee Report

2 major / 2 minor

Summary. The paper claims that mutual information serves as a valid global measure of mixing efficiency in complex flows generated by microswimmers. Using direct numerical computation of flows from a 2D confined squirmer model and solution of the advection-diffusion equation for tracers, the authors report that mixing depends on swimmer spatial distribution (suppressed by aggregation, enhanced by disorder), exhibits non-monotonic dependence on the squirmer parameter at fixed dissipation due to a translation-dipolar flow balance, shows pushers outperforming pullers with hydrodynamic interactions, and displays three-stage temporal decay of mutual information (diffusion-dominated, advection-enhanced, and system-size relaxation). This demonstrates applicability of mutual information beyond simplified models and identifies a shear-dispersion trade-off in microswimmer mixing.

Significance. If the numerical results hold, the work provides a concrete demonstration that an information-theoretic mixing metric extends to swimmer-generated flows without parameter fitting, yielding falsifiable predictions such as the optimal finite squirmer parameter and pusher/puller ordering. The approach credits direct computation of flows and advection-diffusion rather than reduced models, and the three-stage decay offers a testable temporal signature. This could inform studies of biological mixing in low-Reynolds-number fluids, though the 2D confinement limits immediate generality.

major comments (2)
  1. [Abstract and model description] Abstract and model section: the central claim that mutual information applies to mixing by real microswimmers and reveals a shear-dispersion trade-off rests on the 2D squirmer model in confinement capturing essential 3D physics (different interaction ranges, out-of-plane flows); no justification, scaling argument, or test is supplied showing why dimensionality reduction preserves the reported non-monotonic β dependence or pusher/puller ordering.
  2. [Numerical methods] Methods/numerical implementation: details on spatial discretization, time-stepping scheme, grid convergence tests, and error control for the advection-diffusion solver are not provided; these are load-bearing for the quantitative claims of non-monotonic dependence on squirmer parameter and the three distinct decay stages of mutual information.
minor comments (2)
  1. [Figures and results] Figure captions and text should explicitly state the number of independent realizations and the precise definition of the squirmer parameter β used in the non-monotonic scans.
  2. [Results on squirmer parameter] The abstract states 'at fixed energy dissipation' but the main text should clarify how dissipation is computed and held constant across β values.

Simulated Author's Rebuttal

2 responses · 0 unresolved

We thank the referee for their thorough review and valuable comments on our manuscript. We address each major comment below and have made revisions to the manuscript where necessary to strengthen the presentation.

read point-by-point responses
  1. Referee: [Abstract and model description] Abstract and model section: the central claim that mutual information applies to mixing by real microswimmers and reveals a shear-dispersion trade-off rests on the 2D squirmer model in confinement capturing essential 3D physics (different interaction ranges, out-of-plane flows); no justification, scaling argument, or test is supplied showing why dimensionality reduction preserves the reported non-monotonic β dependence or pusher/puller ordering.

    Authors: We agree that an explicit justification for the 2D model was missing from the original manuscript. In the revised version, we will add a paragraph in the model section providing a scaling argument: the essential physics of the shear-dispersion trade-off originates from the competition between the translational velocity (which scales with the squirmer parameter β) and the dipolar flow strength, both of which have direct analogs in 3D squirmer models. The non-monotonic dependence and pusher/puller asymmetry arise from this balance and the form of the hydrodynamic interactions, which are qualitatively similar in 2D and 3D for confined geometries. While we acknowledge that 3D simulations would provide further validation, the 2D confined setup is standard for such studies and enables the high-fidelity computations needed to resolve the mutual information accurately. We believe this addition addresses the concern. revision: yes

  2. Referee: [Numerical methods] Methods/numerical implementation: details on spatial discretization, time-stepping scheme, grid convergence tests, and error control for the advection-diffusion solver are not provided; these are load-bearing for the quantitative claims of non-monotonic dependence on squirmer parameter and the three distinct decay stages of mutual information.

    Authors: The referee correctly identifies that the numerical methods section lacks sufficient detail. We will revise the manuscript to include a new subsection under Methods that specifies: the spatial discretization scheme (e.g., finite differences or spectral methods used for the flow and advection-diffusion equations), the time-stepping algorithm (including any stability criteria), the grid resolutions employed, results of convergence tests demonstrating that the mutual information values and decay stages are insensitive to further refinement within the reported precision, and the error control measures (such as tolerance for iterative solvers). These additions will substantiate the quantitative results without altering the findings. revision: yes

Circularity Check

0 steps flagged

Numerical solution of advection-diffusion equation yields mutual information without reduction to inputs by construction

full rationale

The paper's central results follow from explicit numerical computation of squirmer-generated flows followed by direct solution of the advection-diffusion equation for tracers and subsequent evaluation of mutual information. No parameters are fitted within the paper to subsets of its own data, no self-citation supplies a load-bearing uniqueness theorem or ansatz, and the reported non-monotonic dependence and stage-wise decay are outputs of the simulation rather than definitions or renamings of the inputs. The derivation chain is therefore self-contained and independent of the patterns that would produce circularity.

Axiom & Free-Parameter Ledger

1 free parameters · 1 axioms · 0 invented entities

The central claim rests on the standard advection-diffusion description of tracer transport and the validity of the 2D squirmer model for generating flows; the squirmer parameter is explored numerically rather than derived from first principles.

free parameters (1)
  • squirmer parameter
    Controls the relative strength of swimmer translation versus dipolar flow generation; the paper reports an optimal finite value arising from their balance.
axioms (1)
  • domain assumption The advection-diffusion equation accurately describes the transport of tracer particles in the flow field generated by the squirmers.
    Invoked when solving for the spatial distribution of tracers to compute mutual information.

pith-pipeline@v0.9.1-grok · 5741 in / 1440 out tokens · 34771 ms · 2026-07-03T17:57:55.326166+00:00 · methodology

discussion (0)

Sign in with ORCID, Apple, or X to comment. Anyone can read and Pith papers without signing in.

Reference graph

Works this paper leans on

70 extracted references · 70 canonical work pages

  1. [1]

    In two- dimensional incompressible Stokes flow, the velocity field generated by a point forceF= (F x, Fy) applied atx s = (xs, ys) satisfies −µ∇2v+∇p=Fδ(x−x s),(A1) with∇ ·v= 0

    Fundamental singularities We first summarize the far-field singularities used to construct the flow generated by a microswimmer. In two- dimensional incompressible Stokes flow, the velocity field generated by a point forceF= (F x, Fy) applied atx s = (xs, ys) satisfies −µ∇2v+∇p=Fδ(x−x s),(A1) with∇ ·v= 0. LetF=αe, wheree= (e 1, e2) is a unit vector. The f...

  2. [2]

    Perfect-slip boundary conditions and image construction We next construct the flow field of a singularity con- fined in a square domain of side lengthL. The square is bounded by perfect-slip walls, for which vx|x=0,L = 0, ∂vy ∂x x=0,L = 0,(B1) vy|y=0,L = 0, ∂vx ∂y y=0,L = 0.(B2) These conditions enforce zero normal velocity while al- lowing tangential sli...

  3. [3]

    Let Nbe the number of grid points along one side of the original square

    Discrete Fourier representation of the confined Stokeslet We solve the image problem using a discrete Fourier transform on the doubled periodic domain 2L×2L. Let Nbe the number of grid points along one side of the original square. One period of the image lattice contains NT = 2(N−1) (B4) grid points in each direction, and the real-space grid spac- ing is ...

  4. [4]

    The first mode is associated with a source-dipole contribution, while the second mode gives the stresslet contribution

    Relation to the two-mode squirmer model The two-dimensional squirmer is described by the tan- gential surface slip velocity vs ϕ(a, ϕ) = ∞X n=1 Bn sin(nϕ).(C1) In this work, only the first two modes are retained. The first mode is associated with a source-dipole contribution, while the second mode gives the stresslet contribution. In terms of the confined...

  5. [5]

    Confined stresslet Taking the derivative of Eq. (B13) with respect tox 0, the confined stresslet is v(c) B2(x) =v B2 mn =−B 2a e· ∇ 0G(c) = 1 N 2 T NT /2−1X k,s=−NT /2 eik·x˜vB2 ks , (C5) where ˜vB2 ks =4πB2ai |k|2h2 (e1, e2)(k·e)e −ik·xs + (−e1, e2)(k2 ·e)e −ik2·xs + (e1,−e 2)(k3 ·e)e −ik3·xs + (−e1,−e 2)(k4 ·e)e −ik4·xs I− kk |k|2 . (C6)

  6. [6]

    Confined source dipole The confined source dipole is v(c) B1(x) =v B1 mn =− a2B1 2 1 2 ∇2 0G(c) = 1 N 2 T NT /2−1X k,s=−NT /2 eik·x˜vB1 ks , (C7) where ˜vB1 ks = πB1a2 h2 (e1, e2)e−ik·xs + (−e1, e2)e−ik2·xs + (e1,−e 2)e−ik3·xs + (−e1,−e 2)e−ik4·xs I− kk |k|2 . (C8) 13

  7. [7]

    We therefore introduce exponential spectral regularization, ˜vks → ˜vkse−ε|k|/π,(C9) whereεis a short-distance regularization length

    Regularized confined squirmer flow The singularity at the swimmer position and the slow decay of the Fourier coefficients can lead to numerical in- stabilities when solving the Fokker–Planck equation. We therefore introduce exponential spectral regularization, ˜vks → ˜vkse−ε|k|/π,(C9) whereεis a short-distance regularization length. The regularized stress...

  8. [8]

    Swimmer trajectories without hydrodynamic interactions In the absence of hydrodynamic interactions, swim- mers translate with the free-space squirmer velocity as shown in Eq. (6). The swimmer moves in a straight line until it reaches a wall, where its swimming direction is re- flected geometrically: the normal component ofechanges sign, while the tangenti...

  9. [9]

    J. P. Heller, An unmixing demonstration, Am. J. Phys 28, 348 (1960)

  10. [10]

    Arrieta, J

    J. Arrieta, J. H. E. Cartwright, E. Gouillart, N. Piro, O. Piro, and I. Tuval, Geometric mixing, Phil. Trans. R. Soc. A.378, 20200168 (2020)

  11. [11]

    Golestanian, J

    R. Golestanian, J. M. Yeomans, and N. Uchida, Hydro- dynamic synchronization at low Reynolds number, Soft Matter7, 3074 (2011)

  12. [12]

    Tang and R

    E. Tang and R. Golestanian, Quantifying configurational information for a stochastic particle in a flow-field, New J. Phys.22, 083060 (2020)

  13. [13]

    Villermaux, Mixing versus stirring, Annu

    E. Villermaux, Mixing versus stirring, Annu. Rev. Fluid Mech.51, 245 (2019)

  14. [14]

    Y. Shi, A. Vilfan, and R. Golestanian, Mutual informa- tion as a measure of mixing efficiency in viscous fluids, Phys. Rev. Research6, L022050 (2024)

  15. [15]

    P. V. Danckwerts, The definition and measurement of some characteristics of mixtures, Appl. Sci. Res.3, 279 (1952)

  16. [16]

    J. L. Thiffeault, Using multiscale norms to quantify mix- ing and transport, Nonlinearity25, R1 (2012)

  17. [17]

    Mathew, I

    G. Mathew, I. Mezi´ c, and L. Petzold, A multiscale mea- sure for mixing, Physica D211, 23 (2005)

  18. [18]

    Meunier and E

    P. Meunier and E. Villermaux, The diffuselet concept for scalar mixing, J. Fluid Mech.951, A33 (2022)

  19. [19]

    Y. K. Tsang, T. M. Antonsen, and E. Ott, Exponential decay of chaotically advected passive scalars in the zero diffusivity limit, Phys. Rev. E71, 066301 (2005)

  20. [20]

    Cocconi, Y

    L. Cocconi, Y. Shi, and A. Vilfan, Information-optimal mixing at low Reynolds number, Phys. Rev. Lett.135, 037101 (2025). 14

  21. [21]

    H. Aref, J. R. Blake, M. Budiˇ si´ c, S. S. S. Cardoso, J. H. E. Cartwright, H. J. H. Clercx, K. El Omari, U. Feudel, R. Golestanian, E. Gouillart, G. F. van Hei- jst, T. S. Krasnopolskaya, Y. Le Guer, R. S. MacKay, V. V. Meleshko, G. Metcalfe, I. Mezi´ c, A. P. S. de Moura, O. Piro, M. F. M. Speetjens, R. Sturman, J. L. Thiffeault, and I. Tuval, Frontier...

  22. [22]

    O. H. Shapiro, V. I. Fernandez, M. Garren, J. S. Guasto, F. P. Debaillon-Vesque, E. Kramarsky-Winter, A. Vardi, and R. Stocker, Vortical ciliary flows actively enhance mass transport in reef corals, Proc. Natl. Acad. Sci. U.S.A.111, 13391 (2014)

  23. [23]

    Gilpin, V

    W. Gilpin, V. N. Prakash, and M. Prakash, Vortex arrays and ciliary tangles underlie the feeding–swimming trade- off in starfish larvae, Nat. Phys.13, 380 (2017)

  24. [24]

    C. J. Campbell and B. A. Grzybowski, Microfluidic mix- ers: from microfabricated to self-assembling devices, Phil. Trans. R. Soc. A.362, 1069 (2004)

  25. [25]

    R. O. Grigoriev, M. F. Schatz, and V. Sharma, Chaotic mixing in microdroplets, Lab. Chip6, 1369 (2006)

  26. [26]

    D. J. Pine, J. P. Gollub, J. F. Brady, and A. M. Leshan- sky, Chaos and threshold for irreversibility in sheared sus- pensions, Nature438, 997 (2005)

  27. [27]

    A. D. Stroock, S. K. W. Dertinger, A. Ajdari, I. Mezi´ c, H. A. Stone, and G. M. Whitesides, Chaotic mixer for microchannels, Science295, 647 (2002)

  28. [28]

    Y. Ding, J. C. Nawroth, M. J. McFall-Ngai, and E. Kanso, Mixing and transport by ciliary carpets: a numerical study, J. Fluid Mech.743, 124 (2014)

  29. [29]

    Jonas, E

    S. Jonas, E. Zhou, E. Deniz, B. Huang, K. Chandrasek- era, D. Bhattacharya, Y. Wu, R. Fan, T. M. Deserno, M. K. Khokha, and M. A. Choma, A novel approach to quantifying ciliary physiology: microfluidic mixing driven by a ciliated biological surface, Lab. Chip13, 4160 (2013)

  30. [30]

    J. C. Nawroth, H. Guo, E. Koch, E. A. C. Heath- Heckman, J. C. Hermanson, E. G. Ruby, J. O. Dabiri, E. Kanso, and M. McFall-Ngai, Motile cilia create fluid- mechanical microhabitats for the active recruitment of the host microbiome, Proc. Natl. Acad. Sci. U.S.A.114, 9510 (2017)

  31. [31]

    A. R. Shields, B. L. Fiser, B. A. Evans, M. R. Falvo, S. Washburn, and R. Superfine, Biomimetic cilia ar- rays generate simultaneous pumping and mixing regimes, Proc. Natl. Acad. Sci. U.S.A.107, 15670 (2010)

  32. [32]

    Rahbar, L

    M. Rahbar, L. Shannon, and B. L. Gray, Microfluidic ac- tive mixers employing ultra-high aspect-ratio rare-earth magnetic nano-composite polymer artificial cilia, J. Mi- cromech. Microeng.24, 025003 (2014)

  33. [33]

    Supatto, S

    W. Supatto, S. E. Fraser, and J. Vermot, An all-optical approach for probing microscopic flows in living embryos, Biophys. J.95, L29 (2008)

  34. [34]

    S. A. Selvan, C. O. Pacherres, M. K¨ uhl, A. Butler, M. S. Dhillon, L. L. Blackall, P. W. Duck, D. Pihler-Puzovi´ c, and D. R. Brumley, Unravelling three-dimensional active transport by ciliary arrays on coral surfaces, PRX Life4, 023020 (2026)

  35. [35]

    Uchida and R

    N. Uchida and R. Golestanian, Synchronization and col- lective dynamics in a carpet of microfluidic rotors, Phys. Rev. Lett.104, 178103 (2010)

  36. [36]

    Kunze, Biologically generated mixing in the ocean, Annu

    E. Kunze, Biologically generated mixing in the ocean, Annu. Rev. Mar. Sci.11, 215 (2019)

  37. [37]

    M. E. Huntley and M. Zhou, Influence of animals on tur- bulence in the sea, Mar. Ecol. Prog. Ser.273, 65 (2004)

  38. [38]

    Kunze, J

    E. Kunze, J. F. Dower, I. Beveridge, R. Dewey, and K. P. Bartlett, Observations of biologically generated turbu- lence in a coastal inlet, Science313, 1768 (2006)

  39. [39]

    Katija and J

    K. Katija and J. O. Dabiri, A viscosity-enhanced mecha- nism for biogenic ocean mixing, Nature460, 624 (2009)

  40. [40]

    Leshansky and L

    A. Leshansky and L. Pismen, Do small swimmers mix the ocean?, Phys. Rev. E82, 025301 (2010)

  41. [41]

    Katija, Biogenic inputs to ocean mixing, J Exp Biol

    K. Katija, Biogenic inputs to ocean mixing, J Exp Biol. 215, 1040 (2012)

  42. [42]

    Wang and A

    S. Wang and A. M. Ardekani, Biogenic mixing induced by intermediate Reynolds number swimming in stratified fluids, Sci. Rep.5, 17448 (2015)

  43. [43]

    V. A. Shaik and G. J. Elfring, Mixing by squirmers in stratified fluids, Phys. Rev. Fluids10, 024102 (2025)

  44. [44]

    X. L. Wu and A. Libchaber, Particle diffusion in a quasi- two-dimensional bacterial bath, Phys. Rev. Lett.84, 3017 (2000)

  45. [45]

    P. T. Underhill, J. P. Hernandez-Ortiz, and M. D. Gra- ham, Diffusion and spatial correlations in suspensions of swimming particles, Phys. Rev. Lett.100, 248101 (2008)

  46. [46]

    J. W. van de Meent, I. Tuval, and R. E. Goldstein, Na- ture’s microfluidic transporter: Rotational cytoplasmic streaming at high P´ eclet numbers, Phys. Rev. Lett.101, 178102 (2008)

  47. [47]

    K. C. Leptos, J. S. Guasto, J. P. Gollub, A. I. Pesci, and R. E. Goldstein, Dynamics of enhanced tracer diffusion in suspensions of swimming eukaryotic microorganisms, Phys. Rev. Lett.103, 198103 (2009)

  48. [48]

    Z. Lin, J. L. Thiffeault, and S. Childress, Stirring by squirmers, J. Fluid Mech.669, 167 (2011)

  49. [49]

    I. M. Zaid, J. Dunkel, and J. M. Yeomans, L´ evy fluc- tuations and mixing in dilute suspensions of algae and bacteria, J. R. Soc. Interface8, 1314 (2011)

  50. [50]

    D. O. Pushkin and J. M. Yeomans, Fluid mixing by curved trajectories of microswimmers, Phys. Rev. Lett. 111, 188101 (2013)

  51. [51]

    D. O. Pushkin and J. M. Yeomans, Stirring by swimmers in confined microenvironments, J. Stat. Mech. , P04030 (2014)

  52. [52]

    J. L. Thiffeault, Distribution of particle displacements due to swimming microorganisms, Phys. Rev. E92, 023023 (2015)

  53. [53]

    Mueller and J

    P. Mueller and J. L. Thiffeault, Fluid transport and mix- ing by an unsteady microswimmer, Phys. Rev. Fluids2, 013103 (2017)

  54. [54]

    Ortlieb, S

    L. Ortlieb, S. Rafa¨ ı, P. Peyla, C. Wagner, and T. John, Statistics of colloidal suspensions stirred by microswim- mers, Phys. Rev. Lett.122, 148101 (2019)

  55. [55]

    Ishikawa, Transport phenomena in microswimmer sus- pensions: migration, collective motion, diffusion and rhe- ology, J

    T. Ishikawa, Transport phenomena in microswimmer sus- pensions: migration, collective motion, diffusion and rhe- ology, J. Fluid Mech.1016, P1 (2025)

  56. [56]

    M. J. Lighthill, On the squirming motion of nearly spher- ical deformable bodies through liquids at very small Reynolds numbers, Commun. Pure Appl. Maths5, 109 (1952)

  57. [57]

    J. R. Blake, A spherical envelope approach to ciliary propulsion, J. Fluid Mech.46, 199 (1971)

  58. [58]

    T. J. Pedley, Spherical squirmers: models for swimming micro-organisms, IMA J. Appl. Maths81, 488 (2016)

  59. [59]

    Wang and A

    S. Wang and A. M. Ardekani, Swimming of a model cili- ate near an air-liquid interface, Phys. Rev. E87, 063010 (2013). 15

  60. [60]

    N. G. Chisholm, D. Legendre, E. Lauga, and A. S. Khair, A squirmer across Reynolds numbers, J. Fluid Mech. 796, 233 (2016)

  61. [61]

    J. R. Blake, Self propulsion due to oscillations on the surface of a cylinder at low Reynolds number, Bull. Aust. Math. Soc.5, 255 (1971)

  62. [62]

    S. E. Spagnolie and E. Lauga, Hydrodynamics of self- propulsion near a boundary: predictions and accuracy of far-field approximations, J. Fluid Mech.700, 105 (2012)

  63. [63]

    Ahana and S

    P. Ahana and S. P. Thampi, Confinement induced tra- jectory of a squirmer in a two dimensional channel, Fluid Dyn. Res.51, 065504 (2019)

  64. [64]

    Nasouri, A

    B. Nasouri, A. Vilfan, and R. Golestanian, Minimum dis- sipation theorem for microswimmers, Phys. Rev. Lett. 126, 034503 (2021)

  65. [65]

    T. M. Cover and J. A. Thomas,Elements of Information Theory, 2nd ed. (John Wiley and Sons, Ltd, 2005)

  66. [66]

    See Supplemental Material at [URL will be inserted by publisher] for movies of the time evolution of the proba- bility density in Fig 2

  67. [67]

    Alarc´ on and I

    F. Alarc´ on and I. Pagonabarraga, Spontaneous aggrega- tion and global polar ordering in squirmer suspensions, J. Mol. Liq.185, 56 (2013)

  68. [68]

    Oyama, J

    N. Oyama, J. J. Molina, and R. Yamamoto, Purely hy- drodynamic origin for swarming of swimming particles, Phys. Rev. E93, 043114 (2016)

  69. [69]

    A. W. Zantop and H. Stark, Emergent collective dynam- ics of pusher and puller squirmer rods: swarming, clus- tering, and turbulence, Soft Matter18, 6179 (2022)

  70. [70]

    Golestanian, Hydrodynamically consistent many- body Harada-Sasa relation, Phys

    R. Golestanian, Hydrodynamically consistent many- body Harada-Sasa relation, Phys. Rev. Lett.134, 207101 (2025)