In their search for metabolic resources microbes swim through viscous environments that present physical anisotropies, including steric obstacles across a wide range of sizes. Hydrodynamic forces are known to significantly alter swimmer trajectories near flat and low-curvature surfaces. In this work, we imaged hundreds-of-thousands of high-curvature scattering interactions between swimming bacteria and micro-fabricated pillars with radii from ~1 to ~10 cell lengths. As a function of impact parameter, cell-pillar interactions produced distinct chiral distributions for scattering angle -- including unexpected 'counter-rotator' trajectories -- well-described by a sterics-only model. Our data and model suggest that alteration of swimmer trajectories is subject to distinct mechanisms when interacting with objects of different size; primarily steric for objects below ~10 cell lengths and requiring incorporation of hydrodynamics at larger scales. These alterations in trajectory impact swim dynamics and may affect microbial populations in ways that depend on the shape and placement of obstacles within an environment.
We describe the consequences of time reversal invariance of the Stokes' equations for the hydrodynamic scattering of two low Reynolds number swimmers. For swimmers that are related to each other by a time reversal transformation this leads to the striking result that the angle between the two swimmers is preserved by the scattering. The result is illustrated for the particular case of a linked-sphere model swimmer. For more general pairs of swimmers, not related to each other by time reversal, we find hydrodynamic scattering can alter the angle between their trajectories by several tens of deg…
We propose a very simple one-dimensional swimmer consisting of three spheres that are linked by rigid rods whose lengths can change between two values. With a periodic motion in a non-reciprocal fashion, which breaks the time-reversal symmetry as well as the translational symmetry, we show that the model device can swim at low Reynolds number. This model system could be used in constructing molecular-size machines.
Here we introduce a two-dimensional (2D) low-Reynolds swimmer and discuss the motion of the swimmer both in noise-free and stochastic regimes. Three spheres, linked by extensible arms, in a plane form the triangle body of micro-swimmer. Expansion and contraction of two-stated linkers in appropriate order causes both translational and rotational movement of the swimmer. It is shown that the motion of the swimmer could be controlled from a rotor to a directed linear swimmer depend on the sequence of the linkers activities. A few amount of noise in the the rhythm of the cycles introduces an inter…
As technological advances allow us to fabricate smaller autonomous self-propelled devices, it is clear that at some point directed propulsion could not come from pre-specified deterministic periodic deformation of the swimmer's body and we need to develop strategies to extract a net directed motion from a series of random transitions in the conformation space of the swimmer. We present a theoretical formulation to describe the "stochastic motor" that drives the motion of low Reynolds number swimmers based on this concept, and use it to study the propulsion of a simple low Reynolds number swimm…
The simple model of a low Reynolds number swimmer made from three spheres that are connected by two arms is considered in its general form and analyzed. The swimming velocity, force--velocity response, power consumption, and efficiency of the swimmer are calculated both for general deformations and also for specific model prescriptions. The role of noise and coherence in the stroke cycle is also discussed.
Swimming and pumping at low Reynolds numbers are subject to the "Scallop theorem", which states that there will be no net fluid flow for time reversible motions. Living organisms such as bacteria and cells are subject to this constraint, and so are existing and future artificial "nano-bots" or microfluidic pumps. We study a very simple mechanism to induce fluid pumping, based on the forced motion of three colloidal beads through a cycle that breaks time-reversal symmetry. Optical tweezers are used to vary the inter-bead distance. This model is inspired by a strut-based theoretical swimmer prop…
We employ three numerical methods to explore the motion of low Reynolds number swimmers, modeling the hydrodynamic interactions by means of the Oseen tensor approximation, lattice Boltzmann simulations and multiparticle collision dynamics. By applying the methods to a three bead linear swimmer, for which exact results are known, we are able to compare and assess the effectiveness of the different approaches. We then propose a new class of low Reynolds number swimmers, generalized three bead swimmers that can change both the length of their arms and the angle between them. Hence we suggest a de…
In the limit of zero Reynolds number (Re), swimmers propel themselves exploiting a series of non-reciprocal body motions. For an artificial swimmer, a proper selection of the power source is required to drive its motion, in cooperation with its geometric and mechanical properties. Although various external fields (magnetic, acoustic, optical, etc.) have been introduced, electric fields are rarely utilized to actuate such swimmers experimentally in unbounded space. Here we use uniform and static electric fields to demonstrate locomotion of a bi-flagellated sphere at low Re via Quincke rotation.…
A recently introduced model for an autonomous swimmer at low Reynolds number that is comprised of three spheres connected by two arms is considered when one of the spheres has a large radius. The Stokes hydrodynamic flow associated with the swimming strokes and net motion of this system can be studied analytically using the Stokes Green's function of a point force in front of a sphere of arbitrary radius $R$ provided by Oseen. The swimming velocity is calculated, and shown to scale as $1/R^3$ with the radius of the sphere.
Swimming in circles occurs in a variety of situations at low Reynolds number. Here we propose a simple model for a swimmer that undergoes circular motion, generalising the model of a linear swimmer proposed by Najafi and Golestanian (Phys. Rev. E 69, 062901 (2004)). Our model consists of three solid spheres arranged in a triangular configuration, joined by two links of time-dependent length. For small strokes, we discuss the motion of the swimmer as a function of the separation angle between its links. We find that swimmers describe either clockwise or anticlockwise circular motion depending o…
We consider a swimmer consisting of a collinear assembly of three spheres connected by two slender rods. This swimmer can propel itself forward by varying the lengths of the rods in a way that is not invariant under time reversal. Although any non-reciprocal strokes of the arms can lead to a net displacement, the energetic efficiency of the swimmer is strongly dependent on the details and sequences of these strokes, and also the sizes of the spheres. We define the efficiency of the swimmer using Lighthill's criterion, i.e., the power that is needed to pull the swimmer by an external force at a…
We computationally studied the pair interactions and collective behavior of asymmetric, dumbbell swimmers over a range of intermediate Reynolds numbers and initial configurations. Depending on the initial positions and the Re, we found that two swimmers either repelled and swum away from one another or assembled one of four stable pairs: in-line and in-tandem, both parallel and anti-parallel. When in these stable pairs, swimmers were coordinated, swum together, and generated fluid flows as one. We compared the stable pairs' speeds, swim direction and fluid flows to those of the single swimmer.…
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…
Metachronal swimming, the sequential beating of limbs with a small phase lag, is observed in many organisms at various scales, but has been studied mostly in the limits of high or low Reynolds numbers. Motivated by the swimming of brine shrimp, a mesoscale organism that operates at intermediate Reynolds numbers, we computationally studied a simple nonreciprocal 2D model that performs metachronal swimming. Our swimmer is composed of two pairs of paddles beating with a phase difference that are symmetrically attached to the sides of a flat body. We numerically solved the Navier-Stokes equations …
Small organisms (e.g., bacteria) and artificial microswimmers move due to a combination of active swimming and passive Brownian motion. Considering a simplified linear three-sphere swimmer, we study how the swimmer size regulates the interplay between self-driven and diffusive behavior at low Reynolds number. Starting from the Kirkwood-Smoluchowski equation and its corresponding Langevin equation, we derive formulas for the orientation correlation time, the mean velocity and the mean square displacement in three space dimensions. The validity of the analytical results is illustrated through nu…
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…
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…
Achieving macroscopic directed migration of microscale swimmers in a fluid is an important step towards utilizing their autonomous motion. It has been experimentally shown that directed motion can be induced, without any external fields, by certain geometrically asymmetric obstacles due to interaction between their boundaries and the swimmers. In this paper, we propose a kinetic-type model to study swimming and directional migration of microscale bimetallic rods in a periodic array of posts with non-circular cross-sections. Both rod position and orientation are taken into account; rod trapping…
A three-dimensional model of a low-Reynold's swimmer is introduced and analyzed in this paper. This model consists of two large and small spheres connected by two perpendicular thin rods. The geometry of this system is motivated by the microorganisms that use a single tail to swim, the large sphere represents the head of microorganism and the small sphere resembles its tail. Each rod changes its length and orientation in a non-reciprocal manner that effectively propel the system. Translational and rotational velocities of the swimmer are studied for different values of parameters. Our findings…
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 apply the geometric theory of swimming at low Reynolds number to the study of nearly circular swimmers in two-dimensional fluids with non-vanishing Hall, or "odd", viscosity. The Hall viscosity gives an off-diagonal contribution to the fluid stress-tensor, which results in a number of striking effects. In particular, we find that a swimmer whose area is changing will experience a torque proportional to the rate of change of the area, with the constant of proportionality given by the coefficient $η^o$ of odd viscosity. After working out the general theory of swimming in fluids with Hall visc…
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 …
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,…
Bead-based micro-swimmers are promising systems for payload delivery on the micro-scale. However, the principles underlying their optimal design are not yet fully understood. Here we study a simple device consisting of three arbitrarily-shaped beads connected by two springs. We analytically determine the most favorable kinematic parameters for sinusoidal driving, and show how the swimmer changes from being a pusher to a puller. For cargo carrying ellipsoidal beads, we perform geometric optimization under the constraint of a constant total volume or surface area, with the aim of maximizing the …
A stiff one-armed swimmer in glycerine goes nowhere, but if its arm is elastic, exerting a restorative torque proportional to local curvature, the swimmer can go on its way. Considering this happy consequence and the principles of elasticity, we study a hyperdiffusion equation for the shape of the elastica in viscous flow, find solutions for impulsive or oscillatory forcing, and elucidate relevant aspects of propulsion. These results have application in a variety of physical and biological contexts, from dynamic biopolymer bending experiments to instabilities of elastic filaments.
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…
Hydrodynamic interactions are crucial for determining the cooperative behavior of microswimmers at low Reynolds numbers. Here we provide a comprehensive analysis of the scaling and strength of the interactions in the case of a pair of three-sphere swimmers with intrinsic elasticity. Both stroke-based and force-based microswimmers are analyzed using an analytic perturbative approach. Following a detailed analysis of the passive interactions, as well as active translations and rotations, we find that the mapping between the stroke-based and force-based swimmers is only possible in a low driving …
Active dumbbell suspensions constitute one of the simplest model system for collective swimming at low Reynolds number. Generalizing recent work, we derive and analyze stroke-averaged equations of motion that capture the effective hydrodynamic far-field interaction between two oscillating, asymmetric dumbbells in three space dimensions. Time-averaged equations of motion, as those presented in this paper, not only yield a considerable speed-up in numerical simulations, they may also serve as a starting point when deriving continuum equations for the macroscopic dynamics of multi-swimmer suspens…
We present a two dimensional model of hydrodynamic interaction between a circular swimmer and a circular post at low Reynolds number, using a point singularity description of the swimming activity. We derive a nonlinear dynamical system fully describing the motion and discuss the generic features of the phase portrait and typical trajectories for a variety of squirmer modes. Contractile swimmers exhibit stable bound orbits arising from the contrasting nature of monopolar and dipolar squirmer modes, which are robust with respect to swimmer size and the inclusion of higher squirmer modes. The be…
Microorganisms are rarely found in Nature swimming freely in an unbounded fluid. Instead, they typically encounter other organisms, hard walls, or deformable boundaries such as free interfaces or membranes. Hydrodynamic interactions between the swimmer and nearby objects lead to many interesting phenomena, such as changes in swimming speed, tendencies to accumulate or turn, and coordinated flagellar beating. Inspired by this class of problems, we investigate locomotion of microorganisms near deformable boundaries. We calculate the speed of an infinitely long swimmer close to a flexible surface…
We present a theory for the self-propulsion of symmetric, half spherical Marangoni boats (soap or camphor boats) at low Reynolds numbers. Propulsion is generated by release (diffusive emission or dissolution) of water-soluble surfactant molecules, which modulate the air-water interfacial tension. Propulsion either requires asymmetric release or spontaneous symmetry breaking by coupling to advection for a perfectly symmetrical swimmer. We study the diffusion-advection problem for a sphere in Stokes flow analytically and numerically both for constant concentration and constant flux boundary cond…
We here demonstrate the experimental realization of inanimate micro-swimmer complexes showing emergent self-propulsion at low Reynolds number Re < 10^(-4) in quasi 2D colloidal systems. Guided by the substrate, self-organized propulsion occurs for up to 25min with typical velocities of 1-3 μm/s, while none of the involved constituents shows self-propulsion on its own.
Dense suspensions of self-propelled rod-like particles exhibit a fascinating variety of non-equilibrium phenomena. By means of computer simulations of a minimal model for rigid self-propelled colloidal rods with variable shape we explore the generic diagram of emerging states over a large range of rod densities and aspect ratios. The dynamics is studied using a simple numerical scheme for the overdamped noiseless frictional dynamics of a many-body system in which steric forces are dominant over hydrodynamic ones. The different emergent states are identified by various characteristic correlatio…
A simple way to generate propulsion at low Reynolds number is to periodically oscillate a passive flexible filament. Here we present a macroscopic experimental investigation of such a propulsive mechanism. A robotic swimmer is constructed and both tail shape and propulsive force are measured. Filament characteristics and the actuation are varied and resulting data are quantitatively compared with existing linear and nonlinear theories.
The possibility of microscopic swimming by extraction of energy from an external flow is discussed, focusing on the migration of a simple trimer across a linear shear flow. The geometric properties of swimming, together with the possible generalization to the case of a vesicle, are analyzed.The mechanism of energy extraction from the flow appears to be the generalization to a discrete swimmer of the tank-treading regime of a vesicle. The swimmer takes advantage of the external flow by both extracting energy for swimming and "sailing" through it. The migration velocity is found to scale linearl…
The optimal strategy for a microscopic swimmer to migrate across a linear shear flow is discussed. The two cases, in which the swimmer is located at large distance, and in the proximity of a solid wall, are taken into account. It is shown that migration can be achieved by means of a combination of sailing through the flow and swimming, where the swimming strokes are induced by the external flow without need of internal energy sources or external drives. The structural dynamics required for the swimmer to move in the desired direction is discussed and two simple models, based respectively on th…
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…
Self-propelled colloids (swimmers) in confining geometries follow trajectories determined by hydrodynamic interactions with the bounding surfaces. However, typically these interactions are ignored or truncated to lowest order. We demonstrate that higher-order hydrodynamic moments cause rod-like swimmers to follow oscillatory trajectories in quiescent fluid between two parallel plates, using a combination of lattice-Boltzmann simulations and far-field calculations. This behavior occurs even far from the confining walls and does not require lubrication results. We show that a swimmer's hydrodyna…
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…
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…
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.
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…
Helical propulsion is used by many microorganisms to swim in viscous-dominated environments. Their swimming dynamics are relatively well understood, but detailed study of the flow fields and actuation mechanisms are still needed to realize wall effects and hydrodynamic interactions. In this letter, we describe the development of an autonomous swimming robot with a helical tail that operates in the Stokes regime. The device uses a battery-based power system with a miniature motor that imposes a rotational speed to a helical tail. The speed, direction, and activation are controlled electronicall…
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…
By numerical modeling we investigate fluid transport in low-Reynolds-number flow achieved with a special elastic filament or artifical cilium attached to a planar surface. The filament is made of superparamagnetic particles linked together by DNA double strands. An external magnetic field induces dipolar interactions between the beads of the filament which provides a convenient way of actuating the cilium in a well-controlled manner. The filament has recently been used to successfully construct the first artificial micro-swimmer [R. Dreyfus at al., Nature 437, 862 (2005)]. In our numerical stu…
Hysteresis has been observed in a study of the transition between laminar flow and vortex shedding in a quasi-two dimensional system. The system is a vertical, rapidly flowing soap film which is penetrated by a rod oriented perpendicular to the film plane. Our experiments show that the transition from laminar flow to a periodic Kármán vortex street can be hysteretic, i.e. vortices can survive at velocities lower than the velocity needed to generate them.
