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Floating active carpets drive transport and aggregation in aquatic ecosystems

Published online by Cambridge University Press:  23 September 2024

G. Aguayo
Affiliation:
Departamento de Física, Facultad de Ciencias Físicas y Matemáticas, Universidad de Chile, Beauchef 850, Santiago, Chile
A.J.T.M. Mathijssen
Affiliation:
Department of Physics and Astronomy, University of Pennsylvania, Philadelphia, PA 19104-6396, USA
H.N. Ulloa*
Affiliation:
Department of Earth and Environmental Science, University of Pennsylvania, Philadelphia, PA 19104-6316, USA
R. Soto
Affiliation:
Departamento de Física, Facultad de Ciencias Físicas y Matemáticas, Universidad de Chile, Beauchef 850, Santiago, Chile
F. Guzmán-Lastra*
Affiliation:
Departamento de Física, Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, Santiago, Chile
*
Email addresses for correspondence: fguzman@uchile.cl, ulloa@sas.upenn.edu
Email addresses for correspondence: fguzman@uchile.cl, ulloa@sas.upenn.edu

Abstract

Communities of swimming microorganisms often thrive near liquid–air interfaces. We study how such ‘active carpets’ shape their aquatic environment by driving biogenic transport in the water column beneath them. The hydrodynamic stirring that active carpets generate leads to diffusive upward fluxes of nutrients from deeper water layers, and downward fluxes of oxygen and carbon. Combining analytical theory and simulations, we examine the biogenic transport by studying fundamental metrics, including the single and pair diffusivity, the first passage time for particle pair encounters and the rate of particle aggregation. Our findings reveal that the hydrodynamic fluctuations driven by active carpets have a region of influence that reaches orders of magnitude further in distance than the size of the organisms. These non-equilibrium fluctuations lead to a strongly enhanced diffusion of particles, which is anisotropic and space dependent. Fluctuations also facilitate encounters of particle pairs, which we quantify by analysing their velocity pair correlation functions as a function of distance between the particles. We found that the size of the particles plays a crucial role in their encounter rates, with larger particles situated near the active carpet being more favourable for aggregation. Overall, this research broadens our comprehension of aquatic systems out of equilibrium and how biologically driven fluctuations contribute to the transport of fundamental elements in biogeochemical cycles.

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), 2024. Published by Cambridge University Press.
Figure 0

Figure 1. (a) An AC near the liquid–air interface generates flows that stir particles suspended below. (b) Strength of hydrodynamic fluctuations as a function of distance from the AC. Markers represent simulation results and lines represent the theoretical prediction from (2.5b). (c) Single-particle diffusivity in the $z$ direction. Circles are results from simulated mean-squared displacements, the squares are results from simulating the variance following (3.3) and the solid orange line represents the theoretical prediction from (2.5b) and (3.3).

Figure 1

Figure 2. (a) Velocity pair correlation in the planar coordinate, $g_{\rho }$, as a function of horizontal particle separation. Coloured circles represent simulation results for different depths, ranging from $z_0 = 2$ (blue) to $z_0 = 20$ (light green), each separated by $\Delta z_0=2$. Lines represent the best fit to the model from (3.5). Inset: these curves collapse when plotted as $d_0/z_0$. The yellow line is the best fit from (3.5), and the pink dashed line is the semi-analytical result obtained by numerically integrating $g_{\rho }(d_0, z_0)$. (b) Pair diffusivity in the planar coordinate, $D^{p}_{\rho }$, as a function of horizontal particle separation, normalised by the single-particle diffusivity. Markers represent simulation values for different depths, $z_0$. The black dashed line is the theoretical asymptotic value from $D_{i}^{p\infty }$. (c,d) Show the pair correlation and the pair diffusion in the vertical coordinate, $g_z$ and $D^p_{z}$, respectively.

Figure 2

Figure 3. (a) Mean first passage time of particle collisions as a function of the horizontal particle separation, $d_0$, for particles initially located at $z_0 = 3$. Squares represent simulations and the line is the prediction $\tau _{FP} \approx \tilde {d_0}^{2}/4 D_{\rho }^p.$ Inset: histogram of first passage times. The vertical red dashed line is the mean first passage time. (b) Clearance time against particle radius for three depths. Symbols show simulation results, and solid lines show $\tau _c\sim 0.7\tau _{FP}$. (c) Average accumulated number of collisions over time. Markers show averaged simulation results for different particle radii, ranging from $a = 3$ (triangles) to $a = 10$ (squares). The background colours represent different depths, ranging from $z_0 = 3$ (light blue) to $z_0 = 9$ (pink).

Figure 3

Figure 4. Comparison between variances measured when the AC is thick and have planar swimmers (purple markers), when the AC is flat and have tilted swimmers (light-green markers), when the AC is thick and have tilted swimmers (orange markers) and when the AC is flat and have planar swimmers (hollow markers, the manuscript case).