arXiv subjects

Eric Lauga

Publications and source records attributed to Eric Lauga.

At least 37 records · Page 2Linked to original sources

arXiv2024

Biophysical Fluid Dynamics in a Petri Dish

The humble Petri dish is perhaps the simplest setting in which to examine the locomotion of swimming organisms, particularly those whose body size is tens of microns to millimetres. The fluid layer in such a container has a bottom no-slip surface and a stress-free upper boundary. It is of fundamental interest to understand the flow fields produced by the elementary and composite singularities of Stokes flow in this geometry. Building on the few particular cases that have previously been considered in the literature, we study here the image systems for the primary singularities of Stokes flow subject to such boundary conditions - the stokeslet, rotlet, source, rotlet dipole, source dipole and stresslet - paying particular attention to the far-field behavior. In several key situations, the depth-averaged fluid flow is accurately captured by the solution of an associated Brinkman equation whose screening length is proportional to the depth of the fluid layer. The case of hydrodynamic bound states formed by spinning microswimmers near a no-slip surface, discovered first using the alga $Volvox$, is reconsidered in the geometry of a Petri dish, where the power-law attractive interaction between microswimmers acquires unusual exponentially screened oscillations.

cond-mat.soft↗

arXiv2023

Controlling confined collective organisation with taxis

Biased locomotion is a common feature of microorganisms, but little is known about its impact on self-organisation. Inspired by recent experiments showing a transition to large-scale flows, we study theoretically the dynamics of magnetotactic bacteria confined to a drop. We reveal two symmetry-breaking mechanisms (one local chiral and one global achiral) leading to self-organisation into global vortices and a net torque exerted on the drop. The collective behaviour is ultimately controlled by the swimmers' microscopic chirality and, strikingly, the system can exhibit oscillations and memory-like features.

cond-mat.soft↗

arXiv2023

Multiflagellarity leads to the size-independent swimming speed of peritrichous bacteria

To swim through a viscous fluid, a flagellated bacterium must overcome the fluid drag on its body by rotating a flagellum or a bundle of multiple flagella. Because the drag increases with the size of bacteria, it is expected theoretically that the swimming speed of a bacterium inversely correlates with its body length. Nevertheless, despite extensive research, the fundamental size-speed relation of flagellated bacteria remains unclear with different experiments reporting conflicting results. Here, by critically reviewing the existing evidence and synergizing our own experiments of large sample sizes, hydrodynamic modeling and simulations, we demonstrate that the average swimming speed of \textit{Escherichia coli}, a premier model of peritrichous bacteria, is independent of their body length. Our quantitative analysis shows that such a counterintuitive relation is the consequence of the collective flagellar dynamics dictated by the linear correlation between the body length and the number of flagella of bacteria. Notably, our study reveals how bacteria utilize the increasing number of flagella to regulate the flagellar motor load. The collective load sharing among multiple flagella results in a lower load on each flagellar motor and therefore faster flagellar rotation, which compensates for the higher fluid drag on the longer bodies of bacteria. Without this balancing mechanism, the swimming speed of monotrichous bacteria generically decreases with increasing body length, a feature limiting the size variation of the bacteria. Altogether, our study resolves a long-standing controversy over the size-speed relation of flagellated bacteria and provides new insights into the functional benefit of multiflagellarity in bacteria.

physics.flu-dyn↗

arXiv2023

Cortex-driven cytoplasmic flows in elongated cells: fluid mechanics and application to nuclear transport in Drosophila embryos

The Drosophila melanogaster embryo, an elongated multi-nucleated cell, is a classical model system for eukaryotic development and morphogenesis. Recent work has shown that bulk cytoplasmic flows, driven by cortical contractions along the walls of the embryo, enable the uniform spreading of nuclei along the anterior-posterior (AP) axis necessary for proper embryonic development. Here we propose two mathematical models to characterise cytoplasmic flows driven by tangential cortical contractions in elongated cells. Assuming Newtonian fluid flow at low Reynolds number in a spheroidal cell, we first compute the flow field exactly, thereby bypassing the need for numerical computations. We then apply our results to recent experiments on nuclear transport in cell cycles 4-6 of Drosophila embryo development. By fitting the cortical contractions in our model to measurements, we reveal that experimental cortical flows enable near-optimal axial spreading of nuclei. A second mathematical approach, applicable to general elongated cell geometries, exploits a long-wavelength approximation to produce an even simpler solution, with errors below 5% compared to the full model. An application of this long-wavelength result to transport leads to fully analytical solutions for the nuclear concentration that capture the essential physics of the system, including optimal axial spreading of nuclei.

physics.flu-dyn↗

arXiv2023

Eukaryotic swimming cells are shaped by hydrodynamic constraints

Eukaryotic swimming cells such as spermatozoa, algae or protozoa use flagella or cilia to move in viscous fluids. The motion of their flexible appendages in the surrounding fluid induces propulsive forces that balance with the viscous drag on the cells and lead to a directed swimming motion. Here, we use our recently built database of cell motility (BOSO-Micro) to investigate the extent to which the shapes of eukaryotic swimming cells may be optimal from a hydrodynamic standpoint. We first examine the morphology of flexible flagella undergoing waving deformation and show that their amplitude-to-wavelength ratio is near the one predicted theoretically to optimise the propulsive efficiency of active filaments. Next, we consider ciliates, for which locomotion is induced by the collective beating of short cilia covering their surface. We show that the aspect ratio of ciliates are close to the one predicted to minimise the viscous drag of the cell body. Both results strongly suggest a key role played by hydrodynamic constraints, in particular viscous drag, in shaping eukaryotic swimming cells.

cond-mat.soft↗

arXiv2023

Theoretical model of confined thermoviscous flows for artificial cytoplasmic streaming

Recent experiments in cell biology have probed the impact of artificially-induced intracellular flows in the spatiotemporal organisation of cells and organisms. In these experiments, mild dynamical heating via focused infrared light from a laser leads to long-range, thermoviscous flows of the cytoplasm inside a cell, a method popularised in cell biology as FLUCS (focused-light-induced cytoplasmic streaming). Here, we present a fully analytical, theoretical model describing the fluid flow and transport of tracers induced by the laser at all length scales in two-dimensional confinement. The focused light causes a small, local temperature change, which in turn results in a small change in the density and viscosity of the fluid locally. We analytically solve for the instantaneous flow field due to the translation of a heat spot of arbitrary time-dependent amplitude along a scan path. We show that the leading-order instantaneous flow results purely from thermal expansion. It is proportional to the magnitude of the temperature perturbation, with far-field behaviour typically dominated by a source or sink flow and proportional to the rate of change of the heat-spot amplitude. In contrast, and in agreement with experimental measurements, the net displacement of a tracer due to a full scan of the heat spot is quadratic in the heat-spot amplitude, as it results from the interplay of thermal expansion and thermal viscosity changes. The corresponding average velocity of tracers over a scan is a hydrodynamic source dipole in the far field, with direction dependent on the relative importance of thermal expansion and thermal viscosity changes. Our quantitative findings show excellent agreement with recent experimental results and will enable the design of new controlled experiments to establish the physiological role of physical transport processes inside cells.

physics.flu-dyn↗

arXiv2022

Microswimmers in vortices: Dynamics and trapping

Biological and artificial microswimmers often self-propel in external flows of vortical nature; relevant examples include algae in small-scale ocean eddies, spermatozoa in uterine peristaltic flows and bacteria in microfluidic devices. A recent experiment has shown that swimming bacteria in model vortices are expelled from the vortex all the way to a well-defined depletion zone (Sokolov and Aranson (2016) "Rapid expulsion of microswimmers by a vortical flow." Nature Comm. 7, 11114). In this paper, we propose a theoretical model to investigate the dynamics of elongated microswimmers in elementary vortices, namely active particles in two- and three-dimensional rotlets. A deterministic model first reveals the existence of bounded orbits near the centre of the vortex and unbounded orbits elsewhere. We further discover a conserved quantity of motion that allows us to map the phase space according to the type of the orbit (bounded vs unbounded). We next introduce translational and rotational noise into the system. Using a Fokker--Planck formalism, we quantify the quality of trapping near the centre of the vortex by examining the probability of escape and the mean time of escape from the region of deterministically bounded orbits. We finally show how to use these findings to formulate a prediction for the radius of the depletion zone, which compares favourably with the experiments of Sokolov and Aranson (2016).

physics.bio-ph↗

arXiv2022

Instability of an active fluid jet

The breakup of a fluid jet into droplets has long fascinated natural scientists, with early research dating back to the 19th century. Infinitesimal perturbations to a jet grow because of surface tension, which eventually leads to breakup of the jet into droplets (Rayleigh-Plateau instability). Although this classical phenomenon has long been studied, it is not clear how it is modified when the fluid is replaced by an active fluid. In this study, we investigate instabilities of a jet of an active fluid. The active fluid is modelled by a suspension of microswimmers that propel themselves by generating surface tangential velocities, i.e. squirmers. The squirmers are assumed to be bottom-heavy and heavier than the surrounding fluid, so that a downward jet of the active fluid self-assembles under gravity. Hydrodynamic interactions between squirmers are computed using the Stokesian dynamics method, in which near-field hydrodynamics are accurately calculated. We find that jets of active fluids are unstable, with different unstable modes between pullers and pushers. For an active fluid of pullers, the jet breaks up into droplets in a varicose manner reminiscent of a Newtonian fluid. For pushers, on the other hand, the jet buckles and undergoes a waving (sinuous) instability. The physical mechanisms for these two instabilities can be understood by an inspection of the stress fields in the jets and parametric study reveals the importance of hydrodynamic interactions in the instabilities. Although both gravity and bottom-heaviness play an essential role in realizing the downward jet, their influence on the instability is found to be limited. Our findings help reveal new features of the collective properties of active fluids.

physics.flu-dyn↗

arXiv2022

Elastohydrodynamic synchronization of rotating bacterial flagella

To rotate continuously without jamming, the flagellar filaments of bacteria need to be locked in phase. While several models have been proposed for eukaryotic flagella, the synchronization of bacterial flagella is less well understood. Starting from a reduced model of flexible and hydrodynamically-coupled bacterial flagella, we rigorously coarse-grain the equations of motion using the method of multiple scales, and hence show that bacterial flagella generically synchronize to zero phase difference via an elastohydrodynamic mechanism. Remarkably, the far-field rate of synchronization is maximized at an intermediate value of elastic compliance, with surprising implications for bacteria.

cond-mat.soft↗

arXiv2022

Jet-driven viscous locomotion of confined thermoresponsive microgels

We consider the dynamics of micro-sized, asymmetrically-coated thermoresponsive hydrogel ribbons (microgels) under periodic heating and cooling in the confined space between two planar surfaces. As the result of the temperature changes, the volume and thus the shape of the slender microgel change, which lead to repeated cycles of bending and elastic relaxation, and to net locomotion. Small devices designed for biomimetic locomotion need to exploit flows that are not symmetric in time (non-reciprocal) to escape the constraints of the scallop theorem and undergo net motion. Unlike other biological slender swimmers, the non-reciprocal bending of the gel centreline is not sufficient here to explain for the overall swimming motion. We show instead that the swimming of the gel results from the flux of water periodically emanating from (or entering) the gel itself due to its shrinking (or swelling). The associated flows induce viscous stresses that lead to a net propulsive force on the gel. We derive a theoretical model for this hypothesis of jet-driven propulsion, which leads to excellent agreement with our experiments.

cond-mat.soft↗

arXiv2022

Order and Information in the Patterns of Spinning Magnetic Micro-disks at the Air-water Interface

The application of the Shannon entropy to study the relationship between information and structures has yielded insights into molecular and material systems. However, the difficulty in directly observing and manipulating atoms and molecules hampers the ability of these systems to serve as model systems for further exploring the links between information and structures. Here, we use, as a model experimental system, hundreds of spinning magnetic micro-disks self-organizing at the air-water interface to generate various spatiotemporal patterns with varying degrees of orders. Using the neighbor distance as the information-bearing variable, we demonstrate the links among information, structure, and interactions. Most importantly, we establish a direct link between information and structure without using explicit knowledge of interactions. Finally, we show that the Shannon entropy by neighbor distances is a powerful observable in characterizing structural changes. Our findings are relevant for analyzing natural self-organizing systems and for designing collective robots.

cond-mat.soft↗

arXiv2021

Hydrodynamic interactions between a point force and a slender filament

The Green's function of the incompressible Stokes equations, the stokeslet, represents the singular flow due to a point force. Its classical value in an unbounded fluid has been extended near surfaces of various shapes, including flat walls and spheres, and in most cases the presence of a surface leads to an advection flow induced at the location of the point force. In this paper, motivated by the biological transport of cargo along polymeric filaments inside eukaryotic cells, we investigate the reaction flow at the location of the point force due to a rigid slender filament located at a separation distance intermediate between the filament radius and its length (i.e. we compute the advection of the point force induced by the presence of the filament). An asymptotic analysis of the problem reveals that the leading-order approximation for the force distribution along the axis of the filament takes a form analogous to resistive-force theory but with drag coefficients that depend logarithmically on the distance between the point force and the filament. A comparison of our theoretical prediction with boundary element computations show good agreement. We finally briefly extend the model to the case of curved filaments.

physics.flu-dyn↗

arXiv2021

Stabilising viscous extensional flows using Reinforcement Learning

The four-roll mill, wherein four identical cylinders undergo rotation of identical magnitude but alternate signs, was originally proposed by GI Taylor to create local extensional flows and study their ability to deform small liquid drops. Since an extensional flow has an unstable eigendirection, a drop located at the flow stagnation point will have a tendency to escape. This unstable dynamics can however be stabilised using, e.g., a modulation of the rotation rates of the cylinders. Here we use Reinforcement Learning, a branch of Machine Learning devoted to the optimal selection of actions based on cumulative rewards, in order to devise a stabilisation algorithm for the four-roll mill flow. The flow is modelled as the linear superposition of four two-dimensional rotlets and the drop is treated as a rigid spherical particle smaller than all other length scales in the problem. Unlike previous attempts to devise control, we take a probabilistic approach whereby speed adjustments are drawn from a probability density function whose shape is improved over time via a form of gradient ascent know as Actor-Critic method. With enough training, our algorithm is able to precisely control the drop and keep it close to the stagnation point for as long as needed. We explore the impact of the physical and learning parameters on the effectiveness of the control and demonstrate the robustness of the algorithm against thermal noise. We finally show that Reinforcement Learning can provide a control algorithm effective for all initial positions and that can be adapted to limit the magnitude of the flow extension near the position of the drop.

physics.flu-dyn↗

arXiv2021

Purely viscous acoustic propulsion of bimetallic rods

Synthetic microswimmers offer models for cell motility and their tunability makes them promising candidates for biomedical applications. Here we measure the acoustic propulsion of bimetallic micro-rods that, when trapped at the nodal plane of a MHz acoustic resonator, swim with speeds of up to 300 microns per second. While past acoustic streaming models predict speeds that are more than one order of magnitude smaller than our measurements, we demonstrate that the acoustic locomotion of the rods is driven by a viscous, non-reciprocal mechanism relying on shape anisotropy akin to that used by swimming cells and that reproduces our data with no adjustable parameters.

cond-mat.soft↗

arXiv2021

Microswimming in viscoelastic fluids

The locomotion of microorganisms and spermatozoa in complex viscoelastic fluids is of critical importance in many biological processes such as fertilization, infection, and biofilm formation. Depending on their propulsion mechanisms, microswimmers display various responses to a complex fluid environment: increasing or decreasing their swimming speed and efficiency, modifying their propulsion kinematics and swimming gaits, and experiencing different hydrodynamic interactions with their surroundings. In this article, we review the fundamental physics of locomotion of biological and synthetic microswimmers in complex viscoelastic fluids. Starting from a continuum framework, we describe the main theoretical approaches developed to model microswimming in viscoelastic fluids, which typically rely on asymptotically small dimensionless parameters. We then summarise recent progress on the mobility of single cells propelled by cilia, waving flagella and rotating helical flagella in unbounded viscoelastic fluids. We next briefly discuss the impact of other physical factors, including the micro-scale heterogeneity of complex biological fluids, the role of Brownian fluctuations of the microswimmers, the effect of polymer entanglement and the influence of shear-thinning viscosity. In particular, for solutions of long polymer chains whose sizes are comparable to the radius of flagella, continuum models cannot be used and instead Brownian Dynamics for the polymers can predict the swimming dynamics. Finally, we discuss the effect of viscoelasticity on the dynamics of microswimmers in the presence of surfaces or external flows and its impact on collective cellular behavior.

cond-mat.soft↗

arXiv2021

Rechargeable self-assembled droplet microswimmers driven by surface phase transitions

The design of artificial microswimmers is often inspired by the strategies of natural microorganisms. Many of these creatures exploit the fact that elasticity breaks the time-reversal symmetry of motion at low Reynolds numbers, but this principle has been notably absent from model systems of active, self-propelled microswimmers. Here we introduce a class of microswimmer that spontaneously self-assembles and swims without using external forces, driven instead by surface phase transitions induced by temperature variations. The swimmers are made from alkane droplets dispersed in aqueous surfactant solution, which start to self-propel upon cooling, pushed by rapidly growing thin elastic tails. When heated, the same droplets recharge by retracting their tails, swimming for up to tens of minutes in each cycle. Thermal oscillations of approximately 5 degrees Celsius induce the swimmers to harness heat from the environment and recharge multiple times. We develop a detailed elastohydrodynamic model of these processes and highlight the molecular mechanisms involved. The system offers a convenient platform for examining symmetry breaking in the motion of swimmers exploiting flagellar elasticity. The mild conditions and biocompatible media render these microswimmers potential probes for studying biological propulsion and interactions between artificial and biological swimmers.

cond-mat.soft↗

arXiv2021

Active particles in linear viscoelastic fluids and the scallop theorem

We derive a general formula for the inertialess dynamics of active particles in linear viscoelastic fluids by means of a modified reciprocal theorem. We then demonstrate that force-free active particles in Maxwell-like linear viscoelastic fluids with no retardation have exactly the same dynamics as in Newtonian fluids. In contrast, active particles in Jeffreys-like fluids with retardation can display markedly different dynamics, including net motion under reciprocal actuation thereby breaking the scallop theorem.

physics.flu-dyn↗

arXiv2021

Asymptotic theory of hydrodynamic interactions between slender filaments

Hydrodynamic interactions (HIs) are important in biophysics research because they influence both the collective and the individual behaviour of microorganisms and self-propelled particles. For instance, HIs at the micro-swimmer level determine the attraction or repulsion between individuals, and hence their collective behaviour. Meanwhile, HIs between swimming appendages (e.g. cilia and flagella) influence the emergence of swimming gaits, synchronised bundles and metachronal waves. In this study, we address the issue of HIs between slender filaments separated by a distance larger than their contour length (d>L) by means of asymptotic calculations and numerical simulations. We first derive analytical expressions for the extended resistance matrix of two arbitrarily-shaped rigid filaments as a series expansion in inverse powers of d/L>1. The coefficients in our asymptotic series expansion are then evaluated using two well-established methods for slender filaments, resistive-force theory (RFT) and slender-body theory (SBT), and our asymptotic theory is verified using numerical simulations based on SBT for the case of two parallel helices. The theory captures the qualitative features of the interactions in the regime d/L>1, which opens the path to a deeper physical understanding of hydrodynamically governed phenomena such as inter-filament synchronisation and multiflagellar propulsion. To demonstrate the usefulness of our results, we next apply our theory to the case of two helices rotating side-by-side, where we quantify the dependence of all forces and torques on the distance and phase difference between them. Using our understanding of pairwise HIs, we then provide physical intuition for the case of a circular array of rotating helices. Our theoretical results will be useful for the study of HIs between bacterial flagella, nodal cilia, and slender microswimmers.

physics.flu-dyn↗