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Interplay between Brownian and hydrodynamic tracer diffusion in suspensions of swimming microorganisms

Published online by Cambridge University Press:  31 October 2023

Henrik Nordanger
Affiliation:
Division of Physical Chemistry, Lund University, Box 124, S-221 00 Lund, Sweden
Alexander Morozov
Affiliation:
SUPA, School of Physics and Astronomy, The University of Edinburgh, James Clerk Maxwell Building, Peter Guthrie Tait Road, Edinburgh EH9 3FD, UK
Joakim Stenhammar*
Affiliation:
Division of Physical Chemistry, Lund University, Box 124, S-221 00 Lund, Sweden
*
Email address for correspondence: joakim.stenhammar@fkem1.lu.se

Abstract

The general problem of tracer diffusion in non-equilibrium baths is important in a wide range of systems, from the cellular level to geographical length scales. In this paper, we revisit the archetypical example of such a system: a collection of small passive particles immersed in a dilute suspension of non-interacting dipolar microswimmers, representing bacteria or algae. In particular, we consider the interplay between thermal (Brownian) diffusion and hydrodynamic (active) diffusion due to the persistent advection of tracers by microswimmer flow fields. Previously, it has been argued that even a moderate amount of Brownian diffusion is sufficient to significantly reduce the persistence time of tracer advection, leading to a significantly reduced value of the effective active diffusion coefficient $D_A$ compared to the non-Brownian case. Here, we show by large-scale simulations and kinetic theory that this effect is in fact practically relevant only for microswimmers that effectively remain stationary while still stirring up the surrounding fluid – so-called shakers. In contrast, for moderate and high values of the swimming speed, relevant for biological microswimmer suspensions, the effect of Brownian motion on $D_A$ is negligible, leading to the effects of advection by microswimmers and Brownian motion being additive. This conclusion contrasts with previous results from the literature, and encourages a reinterpretation of recent experimental measurements of $D_A$ for tracer particles of varying size in bacterial suspensions.

Information

Type
JFM Papers
Creative Commons
Creative Common License - CCCreative Common License - BY
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0), which permits unrestricted re-use, distribution and reproduction, provided the original article is properly cited.
Copyright
© The Author(s), 2023. Published by Cambridge University Press.
Figure 0

Figure 1. Tracer trajectories for infinite and finite swimmer paths. (a) Typical trajectory for a non-diffusing tracer advected by a non-tumbling, point-dipole swimmer following an effectively infinite, straight path. (b) Corresponding trajectory terminated due to a tumbling event. Note that, per (1.1), the effective tracer diffusion $D_A$ is independent of tumbling rate $\lambda$ for sufficiently high swimming speeds when averaged over all possible swimmer–tracer configurations, even though the net displacement $\varDelta$ is much larger for the tumbling swimmer. Tracer trajectories were obtained through direct numerical integration using a non-regularised dipolar flow field, as described by Morozov & Marenduzzo (2014).

Figure 1

Figure 2. Effective tracer diffusion in the absence of Brownian motion. Panel (a) shows the $\lambda ^{-1}$ dependence of ${D_A(D_0=0)}$ for shakers with $v_s = 0$, and panel (b) its $v_s$ dependence at constant $\lambda = 10^{-4}$. Symbols denote simulation results, and solid lines show results from (3.5) using $\varepsilon$ as a fitting parameter. Error bars represent one standard deviation as obtained from averaging over four separate runs with different initial conditions. The results are non-dimensionalised in terms of $\kappa$ and $\varepsilon$.

Figure 2

Figure 3. Brownian motion suppresses active diffusion for slow swimming speeds. (a) Values of $D_A$ measured from LB simulations (diamonds) and calculated from (3.9) (solid lines), both expressed in LB units. (b) The same data but expressed in the dimensionless quantities $\xi$ and $Pe$. For very slow swimmers with $L \lesssim 1$, $D_A$ is reduced compared to the non-Brownian value ($\xi = 1$) whenever ${Pe} < 1$, while for faster swimmers, significantly lower values of $Pe$ are necessary to affect $D_A$. The circles for $L = 0$ and 2.4 correspond to the hydrodynamic diffusion of non-Brownian tracers measured in a suspension of Brownian swimmers of the same $D_0$, verifying the statistical equivalence between tracer and swimmer diffusion in the non-interacting limit. Error bars represent one standard deviation as obtained from averaging over four separate runs with different initial conditions. The results in (a) are non-dimensionalised using $\kappa$ and $\varepsilon$.

Figure 3

Figure 4. Brownian motion decorrelates tracer trajectories. (a,b) Fluid velocity autocorrelation $C_T(t)$ in the co-moving tracer frame measured from LB simulations for (a) shakers ($L=0$), and (b) swimmers with $L = 2.4$ at indicated values of $Pe$. The dotted line shows the correlation function $C_U(t)$ of the fluid velocity in the lab frame, demonstrating that the stationary-tracer approximation $C_T(t) \approx C_U(t)$ is excellent in the absence of Brownian tracer diffusion ($Pe \rightarrow \infty$). (c,d) Corresponding lab-frame correlation functions $C_U(t)$ obtained from kinetic theory (3.9) for a suspension of diffusing swimmers, as described in § 3. Throughout, $C_T$ and $C_U$ are non-dimensionalised using $\kappa$ and $\varepsilon$.

Figure 4

Figure 5. Reduction of $D_A$ varies with microswimmer density. (a) Plots of $\xi$ as a function of $Pe$ for shakers ($L=0$) at various densities $n$, as indicated. (b) Plot of $\xi$ as a function of $n$, at a fixed value of ${Pe} = 0.013$. Simulation data are given by the symbols, with error bars obtained from averaging over four separate runs with different initial conditions, while solid lines are computed from (3.9).