We investigate the behavior of a treadmilling microswimmer in a two-dimensional unbounded domain with a semi-infinite no-slip wall. The wall can also be regarded as a probe or pipette inserted into the flow. We solve the governing evolution equations in an analytical form and numerically calculate trajectories of the swimmer for several different initial positions and orientations. We then compute the probability that the treadmilling organism can escape the vicinity of the wall. We find that many trajectories in a 'wedge' around the wall are likely to escape. This suggests that inserting a probe or pipette in a suspension of organism may push away treadmilling swimmers.
We investigate the hydrodynamic interactions between microorganisms swimming at low Reynolds number. By considering simple model swimmers, and combining analytic and numerical approaches, we investigate the time-averaged flow field around a swimmer. At short distances the swimmer behaves like a pump. At large distances the velocity field depends on whether the swimming stroke is invariant under a combined time-reversal and parity transformation. We then consider two swimmers and find that the interaction between them consists of two parts; a dead term, independent of the motion of the second s…
We computationally study the kinematics of a simple model reciprocal swimmer (asymmetric dumbbell) as a function of the Reynolds number (Re) and investigate how the onset and gradual increase of inertia impacts the swimming behavior: a reversal in the swim direction, flow directions, and the swim stroke. We divide the swim stroke into the expansion and compression of the two spheres and relate them to power and recovery strokes. We find that the switch in swim direction also corresponds to a switch in power and recovery strokes. We obtain expressions for the mean swimming velocity by collapsin…
We use the boundary element method to study the low-Reynolds number locomotion of a spherical model microorganism in a circular tube. The swimmer propels itself by tangen- tial or normal surface motion in a tube whose radius is on the order of the swimmer size. Hydrodynamic interactions with the tube walls significantly affect the average swimming speed and power consumption of the model microorganism. In the case of swimming parallel to the tube axis, the locomotion speed is always reduced (resp. increased) for swimmers with tangential (resp. normal) deformation. In all cases, the rate of wor…
The three-sphere swimmer by Najafi and Golestanian is composed of three spheres connected by two arms. The case in which the swimmer can control the lengths of the two arms has been studied in detail. Here we study a variation of the model in which the swimmer's arms are constructed according to Hill's model of muscular contraction. The swimmer is able to control the tension developed in the active components of the arms. The two shape parameters and the tensions acting on the two arms are then obtained by solving a system of ordinary differential equations. We study the qualitative properties…
Inspired by recent experiments using synthetic microswimmers to manipulate droplets, we investigate the low-Reynolds-number locomotion of a model swimmer (a spherical squirmer) encapsulated inside a droplet of comparable size in another viscous fluid. Meditated solely by hydrodynamic interactions, the encaged swimmer is seen to be able to propel the droplet, and in some situations both remain in a stable co-swimming state. The problem is tackled using both an exact analytical theory and a numerical implementation based on boundary element method, with a particular focus on the kinematics of th…
We describe experiments and simulations demonstrating the propulsion of a neutrally-buoyant swimmer that consists of a pair of spheres attached by a spring, immersed in a vibrating fluid. The vibration of the fluid induces relative motion of the spheres which, for sufficiently large amplitudes, can lead to motion of the center of mass of the two spheres. We find that the swimming speed obtained from both experiment and simulation agree and collapse onto a single curve if plotted as a function of the streaming Reynolds number, suggesting that the propulsion is related to streaming flows. There …
In this fluid dynamics video, we show that low Reynolds number swimmers, such as Caenorhabditis (C.) elegans, synchronize their gait when swimming in close proximity to maximize utilization of space. Synchronization most likely results from steric hindrance and enhances the propulsive speed only marginally.
Understanding the stochastic dynamics of tracer particles in active fluids is important for identifying the physical properties of flow generating objects such as colloids, bacteria or algae. Here, we study both analytically and numerically the scattering of a tracer particle in different types of time-dependent, hydrodynamic flow fields. Specifically, we compare the tracer motion induced by an externally driven colloid with the one generated by various self-motile, multi-sphere swimmers. Our results suggest that force-free swimmers generically induce loop-shaped tracer trajectories. The speci…
Unsteadiness occurs in the low Reynolds number swimmers' motion while they start from rest or escape from a predator or attack prey. In this paper, we study an unsteady chiral swimmer's behavior, with a prescribed surface slip velocity, in the low Reynold number regime and its response to an external chemical gradient. Using the general solution of unsteady Stokes equation and appropriate boundary conditions, i.e., slip velocity at the body surface and the ambient velocity field at the far-field, we calculate the migration velocity, rotation rate, and the ow field of the swimmer in a closed-fo…
Swimming cells and microorganisms are a critical component of many biological processes. In order to better interpret experimental studies of low Reynolds number swimming, we combine experimental and numerical methods to perform an analysis of the flow-field around the swimming nematode Caenorhabditis elegans. We first use image processing and particle tracking velocimetry to extract the body shape, kinematics, and flow-fields around the nematode. We then construct a three-dimensional model using the experimental swimming kinematics and employ a boundary element method to simulate flow-fields,…
Swimming at a micrometer scale demands particular strategies. Indeed when inertia is negligible as compared to viscous forces (i.e. Reynolds number $Re$ is lower than unity), hydrodynamics equations are reversible in time. To achieve propulsion at low Reynolds number, swimmers must then deform in a way that is not invariant under time reversal. Here, we investigate dispersal properties of self propelled organisms by means of microscopy and cell tracking. Our system of interest is the micro-alga \textit{Chlamydomonas Reinhardtii}, a motile single celled green alga about 10 micrometers in diamet…
We investigate a self-organized swimmer at low Reynolds numbers. The microscopic swimmer is composed of three spheres that are connected by two identical active linker arms. Each linker arm contains molecular motors and elastic elements and can oscillate spontaneously. We find that such a system immersed in a viscous fluid can self-organize into a state of directed swimming. The swimmer provides a simple system to study important aspects of the swimming of micro-organisms.
Recently, the Brownian dynamics of self-propelled (active) rod-like particles was explored to model the motion of colloidal microswimmers, catalytically-driven nanorods, and bacteria. Here, we generalize this description to biaxial particles with arbitrary shape and derive the corresponding Langevin equation for a self-propelled Brownian spinning top. The biaxial swimmer is exposed to a hydrodynamic Stokes friction force at low Reynolds numbers, to fluctuating random forces and torques as well as to an external and an internal (effective) force and torque. The latter quantities control its sel…
The effects of fluid viscosity on the kinematics of a small swimmer at low Reynolds number are investigated in both experiments and in a simple model. The swimmer is the nematode Caenorhabditis elegans, which is an undulating roundworm approximately 1 mm long. Experiments show that the nematode maintains a highly periodic swimming behavior as the fluid viscosity is varied from 1.0 mPa-s to 12 mPa-s. Surprisingly, the nematode's swimming speed (~0.35 mm/s) is nearly insensitive to the range of fluid viscosities investigated here. However, the nematode's beating frequency decreases to an asympto…
To achieve propulsion at low Reynolds number, a swimmer must deform in a way that is not invariant under time-reversal symmetry; this result is known as the scallop theorem. We show here that there is no many-scallop theorem. We demonstrate that two active particles undergoing reciprocal deformations can swim collectively; moreover, polar particles also experience effective long-range interactions. These results are derived for a minimal dimers model, and generalized to more complex geometries on the basis of symmetry and scaling arguments. We explain how such cooperative locomotion can be rea…
We consider the tracer diffusion $D_{rr}$ that arises from the run-and-tumble motion of low Reynolds number swimmers, such as bacteria. Assuming a dilute suspension, where the bacteria move in uncorrelated runs of length $λ$, we obtain an exact expression for $D_{rr}$ for dipolar swimmers in three dimensions, hence explaining the surprising result that this is independent of $λ$. We compare $D_{rr}$ to the contribution to tracer diffusion from entrainment.
Both natural and artificial small-scale swimmers may often self-propel in environments subject to complex geometrical constraints. While most past theoretical work on low-Reynolds number locomotion addressed idealised geometrical situations, not much is known on the motion of swimmers in heterogeneous environments. As a first theoretical model, we investigate numerically the behaviour of a single spherical micro-swimmer located in an infinite, periodic body-centred cubic lattice consisting of rigid inert spheres of the same size as the swimmer. Running a large number of simulations we uncover …
Swimming cells often have to self-propel through fluids displaying non-Newtonian rheology. While past theoretical work seems to indicate that stresses arising from complex fluids should systematically hinder low-Reynolds number locomotion, experimental observations suggest that locomotion enhancement is possible. In this paper we propose a physical mechanism for locomotion enhancement of microscopic swimmers in a complex fluid. It is based on the fact that micro-structured fluids will generically phase-separate near surfaces, leading to the presence of low-viscosity layers which promote slip a…
We experimentally and theoretically study the dynamics of a low-Reynolds number helical swimmer moving across viscosity gradients. Experimentally, a double-layer viscosity is generated by superposing two miscible fluids with similar densities but different dynamic viscosities. A synthetic helical magnetically-driven swimmer is then made to move across the viscosity gradients along four different configurations: either head-first (pusher swimmer) or tail-first (puller), and through either positive (i.e. going from low to high viscosity) or negative viscosity gradients. We observe qualitative di…
Biological organisms swimming at low Reynolds number are often influenced by the presence of rigid boundaries and soft interfaces. In this paper we present an analysis of locomotion near a free surface with surface tension. Using a simplified two-dimensional singularity model, and combining a complex variable approach with conformal mapping techniques, we demonstrate that the deformation of a free surface can be harnessed to produce steady locomotion parallel to the interface. The crucial physical ingredient lies in the nonlinear hydrodynamic coupling between the disturbance flow created by th…
Emergent swimming strategies of a smart three-bead swimmer
Institute for Theoretical Physics, TU Wien（维也纳工业大学理论物理研究所）；Institute of Science and Technology Austria (ISTA)（奥地利科学技术研究所）；Institute of Computer Engineering, TU Wien（维也纳工业大学计算机工程研究所）；Allen Discovery Center, Tufts University（塔夫茨大学艾伦发现中心）
Hydrodynamic interaction strongly influences the collective behavior of the microswimmers. With this work, we study the behavior of two hydrodynamically interacting self-propelled chiral swimmers in the low Reynolds number regime, considering both the near and far-field interactions. We use the chiral squirmer model, a spherically shaped body with non-axisymmetric surface slip velocity, which generalizes the well-known squirmer model. We calculate the lubrication force between the swimmers when they are very close to each other. By varying the slip coefficients and the initial configuration of…
Reciprocal movement cannot be used for locomotion at low-Reynolds number in an infinite fluid or near a rigid surface. Here we show that this limitation is relaxed for a body performing reciprocal motions near a deformable interface. Using physical arguments and scaling relationships, we show that the nonlinearities arising from reciprocal flow-induced interfacial deformation rectify the periodic motion of the swimmer, leading to locomotion. Such a strategy can be used to move toward, away from, and parallel to any deformable interface as long as the length scales involved are smaller than int…
This contribution provides a general framework to use Lagrange multipliers for the simulation of low Reynolds number fiber dynamics based on Bead Models (BM). This formalism provides an efficient method to account for kinematic constraints. We illustrate, with several examples, to which extent the proposed formulation offers a flexible and versatile framework for the quantitative modeling of flexible fibers deformation and rotation in shear flow, the dynamics of actuated filaments and the propulsion of active swimmers. Furthermore, a new contact model called Gears Model is proposed and success…
We study synchronization in bulk suspensions of spherical microswimmers with chiral trajectories using large scale numerics. The model is generic. It corresponds to the lowest order solution of a general model for self-propulsion at low Reynolds numbers, consisting of a nonaxisymmetric rotating source dipole. We show that both purely circular and helical swimmers can spontaneously synchronize their rotation. The synchronized state corresponds to velocity alignment with high orientational order in both the polar and azimuthal directions. Finally, we consider a racemic mixture of helical swimmer…
Obtaining a detailed understanding of the physical interactions between a cell and its environment often requires information about the flow of fluid surrounding the cell. Cells must be able to effectively absorb and discard material in order to survive. Strategies for nutrient acquisition and toxin disposal, which have been evolutionarily selected for their efficacy, should reflect knowledge of the physics underlying this mass transport problem. Motivated by these considerations, in this paper we discuss the results from an undergraduate research project on the advection-diffusion equation at…
Geometric confinements are frequently encountered in soft matter systems and in particular significantly alter the dynamics of swimming microorganisms in viscous media. Surface-related effects on the motility of microswimmers can lead to important consequences in a large number of biological systems, such as biofilm formation, bacterial adhesion and microbial activity. On the basis of low-Reynolds-number hydrodynamics, we explore the state diagram of a three-sphere microswimmer under channel confinement in a slit geometry and fully characterize the swimming behavior and trajectories for neutra…
In several biologically relevant situations, cell locomotion occurs in polymeric fluids with Weissenberg {number} larger than one. Here we present results of three-dimensional numerical simulations for the steady locomotion of a self-propelled body in a model polymeric (Giesekus) fluid at low Reynolds number. Locomotion is driven by steady tangential deformation at the surface of the body (so-called squirming motion). In the case of a spherical squirmer, we show that the swimming velocity is systematically less than that in a Newtonian fluid, with a minimum occurring for Weissenberg numbers of…
We use numerical simulations to address locomotion at zero Reynolds number in viscoelastic (Giesekus) fluids. The swimmers are assumed to be spherical, to self-propel using tangential surface deformation, and the computations are implemented using a finite element method. The emphasis of the study is on the change of the swimming kinematics, energetics, and flow disturbance from Newtonian to viscoelastic, and on the distinction between pusher and puller swimmers. In all cases, the viscoelastic swimming speed is below the Newtonian one, with a minimum obtained for intermediate values of the Wei…
