The hydrodynamics of swimming microorganisms
arXiv:0812.2887 · doi:10.1088/0034-4885/72/9/096601
Abstract
Cell motility in viscous fluids is ubiquitous and affects many biological processes, including reproduction, infection, and the marine life ecosystem. Here we review the biophysical and mechanical principles of locomotion at the small scales relevant to cell swimming (tens of microns and below). The focus is on the fundamental flow physics phenomena occurring in this inertia-less realm, and the emphasis is on the simple physical picture. We review the basic properties of flows at low Reynolds number, paying special attention to aspects most relevant for swimming, such as resistance matrices for solid bodies, flow singularities, and kinematic requirements for net translation. Then we review classical theoretical work on cell motility: early calculations of the speed of a swimmer with prescribed stroke, and the application of resistive-force theory and slender-body theory to flagellar locomotion. After reviewing the physical means by which flagella are actuated, we outline areas of active research, including hydrodynamic interactions, biological locomotion in complex fluids, the design of small-scale artificial swimmers, and the optimization of locomotion strategies.
Review article
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- Stability and dynamics of magnetocapillary interactions
- Simulation of microswimmer hydrodynamics with multiparticle collision dynamics
- Orbital Magnetism of Active Viscoelastic Suspension
- Hydrodynamic interactions hinder transport of flow-driven colloidal particles
- Trading particle shape with fluid symmetry: on the mobility matrix in 3D chiral fluids
- Generalized Three-Sphere Microswimmers
- Weakly nonlinear analysis of pattern formation in active suspensions
- The hydrodynamics of slender swimmers near deformable interfaces
- Self-propulsion in 2D Confinement: Phoretic and Hydrodynamic Interactions
- A general perturbative approach for bead-based microswimmers reveals rich self-propulsion phenomena
- Biopolymer dynamics driven by helical flagella
- Simulating active agents under confinement with Dissipative Particles (hydro)Dynamics
- Shapes of sedimenting soft elastic capsules in a viscous fluid
- Optimal Ciliary Locomotion of Axisymmetric Microswimmers
- Minimal two-sphere model of the generation of fluid flow at low Reynolds numbers
- Helical micropumps near surfaces
- Asymmetric bistability of chiral particle orientation in viscous shear flows
- Effect of fluid inertia on swimming of a sphere in a viscous incompressible fluid
- Purely viscous acoustic propulsion of bimetallic rods
- Self-diffusiophoretic propulsion of a spheroidal particle in a shear-thinning fluid
- Efficient sliding locomotion with isotropic friction
- The role of tumbling in bacterial scattering at convex obstacles
- Diffusiophoretic propulsion of an isotropic active colloidal particle near a finite-sized disk embedded in a planar fluid-fluid interface
- The effects of rapid yawing on simple swimmer models and planar Jeffery's orbits
- Emergent pattern formation of active magnetic suspensions in an external field
- Large scale zigzag pattern emerging from circulating active shakers
- Simulating wet active polymers by multiparticle collision dynamics
- From active stresses and forces to self propulsion of droplets
- The non-Gaussian tops and tails of diffusing boomerangs
- Dipolophoresis in concentrated suspensions of ideally polarizable spheres
- Theory of active particle penetration through a planar elastic membrane
- Swimming of bacterium \textit{Bacillus subtilis} with multiple bundles of flagella
- Cargo Towing by Artificial Swimmers
- Phase coexistence in a monolayer of active particles induced by Marangoni flows
- Mode instabilities and dynamic patterns in a colony of self-propelled surfactant particles covering a thin liquid layer
- Motility and self-organization of gliding Chlamydomonas populations
- Dynamics of flexible fibers in confined shear flows at finite Reynolds numbers
- Formation of a strong negative wake behind a helical swimmer in a viscoelastic fluid
- Pair Dispersion in Dilute Suspension of Active Swimmers
- Hydrodynamic interaction between two elastic microswimmers
- Self-buckling and self-writhing of semi-flexible microorganisms
- Collective behavior of squirmers in thin films
- Periodic Orbits of Active Particles induced by Hydrodynamic Monopoles
- Electrophoresis of metal-dielectric Janus particles with dipolar director symmetry in nematic liquid crystals
- Swinging Motion of Active Deformable Particles in Poiseuille Flow
- Alignment-Induced Self-Organization of Autonomously Steering Microswimmers: Turbulence, Vortices, and Jets
- Collinear swimmer propelling a cargo sphere at low Reynolds number
- Inertia drives concentration-wave turbulence in swimmer suspensions
- Defect turbulence in a dense suspension of polar, active swimmers
- Confinement induced three-dimensional trajectories of microswimmers in rectangular channels
- Swimmer-microrheology
- Snapping elastic disks as microswimmers: swimming at low Reynolds numbers by shape hysteresis
- Micropropulsion and microrheology in complex fluids via symmetry breaking
- A numerical method for suspensions of articulated bodies in viscous flows
- Transition and formation of the torque pattern of undulatory locomotion in resistive force dominated media
- Polarisation of cells and soft objects driven by mechanical interactions: Consequences for migration and chemotaxis
- Stirrers and movers actuated by oscillating fields
- Drag anisotropy of cylindrical solids in fluid-saturated granular beds
- Three-disk microswimmer in a supported fluid membrane
- Unsteady and inertial dynamics of an active particle in a fluid
- Generalized Scallop Theorem for Linear Swimmers
- Nanoconfined catalytic Ångström-size motors
- Two applications of stochastic thermodynamics to hydrodynamics
- Force and torque-free helical tail robot to study low Reynolds number microorganism swimming
- Locomotion of magnetoelastic membranes in viscous fluids
- Enhanced velocity fluctuations in interacting swimmer suspensions
- Rotation of slender swimmers in isotropic-drag media
- Orbits, spirals, and trapped states: Dynamics of a phoretic Janus particle in a radial concentration gradient
- Axisymmetric monopole and dipole flow singularities in proximity of a stationary no-slip plate immersed in a Brinkman fluid
- Dissolution-driven propulsion of floating solids
- Irreversibility and entropy production of a thermally driven micromachine
- Viscous pumping inspired by flexible propulsion
- A coin vibrational motor swimming at low Reynolds number
- Collective dynamics of active dumbbells near a circular obstacle
- The 3D coarse-graining formulation of interacting elastohydrodynamic filaments and multi-body microhydrodynamics
- Stability of sedimenting flexible loops
- Deformations of an active liquid droplet
- Focusing by blocking: repeatedly generating central density peaks in self-propelled particle systems by exploiting diffusive processes
- Efficient sliding locomotion of three-link bodies
- Competing effects of inertia, sheet elasticity, and fluid viscoelasticity on the synchronization of two actuated sheets
- Extensibility enables locomotion under isotropic drag
- Collective stiffening of soft hair assemblies
- Passively parallel regularized stokeslets
- Stealthy Movements and Concealed Swarms of Swimming micro-Robots
- Exotic Topology in Ordinary Soft Matter
- Propulsion via flexible flapping in granular media
- Using Surface-Motions for Locomotion of Microscopic Robots in Viscous Fluids
- Enhancement of microorganism swimming speed in active matter
- Slender body theories for rotating filaments
- Viscoelastic levitation
- Neuroevolution of Decentralized Decision-Making in N-Bead Swimmers Leads to Scalable and Robust Collective Locomotion
- Phoretic flow induced by asymmetric confinement
- Flow patterns and defect dynamics of active nematics under an electric field
- Well-posedness and applications of classical elastohydrodynamics for a swimming filament
- Deformation of a flexible fiber in a viscous flow past an obstacle
- Dynamics of magnetoelastic robots in water-saturated granular beds
- Load response of shape-changing microswimmers scales with their energy efficiency
- Forced microrheology of active colloids
- Measurements and modelling of induced flow in collective vertical migration
- Non-Stokes Drag Coefficient in Single-Particle Electrophoresis: New Insights on a Classical Problem
- Dynamics of Deformable Active Particles under External Flow Field
- Motion of several slender rigid filaments in a Stokes flow
- A stochastic model for directional changes of swimming bacteria
- Anomalous velocity distributions in active Brownian suspensions
- Active-Passive Brownian Particle in Two Dimensions
- Schrodinger dynamics and Berry phase of undulatory locomotion
- Elastic displacements and viscous flows in wedge-shaped geometries with a straight edge: Green's functions for perpendicular forces
- Long-wavelength instabilities in a system of interacting active particles
- Swimming of the midge larva: principles and tricks of locomotion at intermediate Reynolds number
- Theoretical justification and error analysis for slender body theory
- Activated diffusiophoresis
- Structural transition of vortices to nonlinear regimes in a dusty plasma
- A single-layer based numerical method for the slender body boundary value problem
- Ray Optics for Gliders
- Chaos and irreversibility of a flexible filament in periodically-driven Stokes flow
- Shapes and dynamic regimes of a polar active fluid droplet under confinement
- Microscopic motility of isolated E. coli flagella
- Gauging nanoswimmer dynamics via the motion of large bodies
- Bending fluctuations in semiflexible, inextensible, slender filaments in Stokes flow: towards a spectral discretization
- Force Dipole Interactions in Tubular Fluid Membranes
- The effect of axisymmetric confinement on propulsion of a three-sphere microswimmer
- Self-organisation of auto-phoretic suspensions in confined shear flows
- Self-sustained patchy turbulence in shear-thinning active fluids
- Swimming of a deformable slab in a viscous incompressible fluid with inertia
- Optimal run-and-tumble in slit-like confinement
- Effective medium model for a suspension of active swimmers
- Active dipolar spheroids in shear flow and transverse field: Population splitting, cross-stream migration and orientational pinning
- Self-propulsion of an active polar drop
- Propulsion by stiff elastic filaments in viscous fluids
- Tank-treading as a means of propulsion in viscous shear flows
- Cilia beating patterns are not hydrodynamically optimal
- A three-sphere microswimmer in a structured fluid
- Modeling of Sedimentation of Particles near Corrugated Surface by Boundary Singularity Method
- Autonomous elastic microswimmer
- Dynamics of Purcell-type microswimmers with active-elastic joints
- Activity-enhanced shear thinning of flexible linear polar polymers
- Energy-optimal strokes for multi-link microswimmers: Purcell's loops and Taylor's waves reconciled
- Hydrodynamic force on a small squirmer moving with a time-dependent velocity at small Reynolds numbers
- Inertial torque on a squirmer
- Hydrodynamic interactions in anomalous rheology of active suspensions
- Viscotaxis of Beating Flagella at Surfaces
- Hydrodynamic attraction and repulsion between asymmetric rotors
- Machine learning acceleration of simulations of Stokesian suspensions
- Co-regulation of multi-cellular swirling by individual cellular chirality and boundaries
- Trajectory Tracking Using Motion Primitives for the Purcell's Swimmer
- Geometric phase methods with Stokes theorem for a general viscous swimmer
- Active colloidal particles in emulsion droplets: A model system for the cytoplasm
- Diffusion dynamics of an overdamped active ellipsoidal particle in two dimensions
- The zero-shear-rate limiting rheological behaviors of ideally conductive particles suspended in concentrated dispersions under an electric field
- Mechanical inhibition of dissipation in a thermodynamically consistent active solid
- Regularized Stokeslet rings - an efficient method for axisymmetric Stokes flow, with application to the growing pollen tube
- Inertia-driven propulsion of asymmetric spinner-dimers at moderate Reynolds numbers
- Interplay between Brownian and hydrodynamic tracer diffusion in suspensions of swimming microorganisms
- Modeling of chemically active particles at an air-liquid interface
- Flow induced by the rotation of two circular cylinders in a viscous fluid
- Physicochemical Hydrodynamics of Droplets out of Equilibrium: A Perspective Review
- Hermitian and non-Hermitian topology in active matter
- Conditions for the propulsion of a colloid surrounded by a mesoscale phase separation
- Stokes equation in a semi-infinite region: generalization of Lamb solution and applications to Marangoni flows
- Second harmonic generation and vortex shedding by a dipole-quadrupole and a quadrupole-octupole swimmer in a viscous incompressible fluid
- Asymmetric Fluid Flow in Helical Pipes Inspired by Shark Intestines
- Effective and efficient modeling of the hydrodynamics for bacterial flagella
- Energy Dissipation by Metamorphic Micro-Robots in Viscous Fluids
- Effects of Flagellar Morphology on Swimming Performance and Directional Control in Microswimmers
- Micro-swimmer collective dynamics in Brinkman flows
- Viscous tweezers: controlling particle orientation with viscosity
- Swimming efficiency in viscosity gradients
- Swimming at small Reynolds number of a planar assembly of spheres in an incompressible viscous fluid with inertia
- Engineering passive swimmers by shaking liquids
- A low-Reynolds-number treadmilling swimmer near a semi-infinite wall
- Flagellum Pumping Efficacy in Shear-Thinning Viscoelastic Fluids
- Irreversibility and Chaos in Active Particle Suspensions
- Intermittent sliding locomotion of a two-link body
- Scaling law for a buckled elastic filament in a shear flow
- Stationary shapes of deformable particles moving at low Reynolds numbers
- The equilibrium fluctuations of a semiflexible filament cross linked into a network
- Hydrodynamics of a Micro-Hunter: Chemotactic Scenario
- Intracellular coupling modulates biflagellar synchrony
- 2D force constraints in the method of regularized Stokeslets
- Active Lévy Matter: Anomalous Diffusion, Hydrodynamics and Linear Stability
- HTMPC: A heavily templated C++ library for large scale particle-based mesoscale hydrodynamics simulations using multiparticle collision dynamics
- Rheologically tuned modes of collective transport in active viscoelastic films
- Unified description of viscous, viscoelastic, or elastic thin active films on substrates
- Propulsion of a three-sphere micro-robot in a porous medium
- Anisotropic frictional model for an object sliding in a granular media
- Boosting micromachine studies with Stokesian Dynamics
- Fluid transport by active elastic membranes
- A rigorous proof of the scallop theorem and a finite mass effect of a microswimmer
- Experimental determination of the propulsion matrix of the body of helical Magnetospirillum magneticum cells
- Accuracy of slender body theory in approximating force exerted by thin fiber on viscous fluid
- Self-propulsion of a spherical electric or magnetic microbot in a polar viscous fluid
- Instability-driven Oscillations of Elastic Microfilaments
- Influences of transversely-isotropic rheology and translational diffusion on the stability of active suspensions
- Swimming of a sphere in a viscous incompressible fluid with inertia
- Mixing by squirmers in stratified fluids
- Efficient sliding locomotion of three-link bodies with inertia
- Rapidly yawing spheroids in viscous shear flow: Emergent loss of symmetry
- Parametric Shape Optimization of Flagellated Micro-Swimmers Using Bayesian Techniques
- Sedimentation and structure of squirmer suspensions under gravity
- Dynamical Boundary Following and Corner Trapping of Undulating Worms
- Non-reciprocal mixtures in suspension: the role of hydrodynamic interactions
- Motility and rotation of multi-timescale microswimmers in linear background flows
- Micro-swimmer locomotion and hydrodynamics in Brinkman Flows
- A Hamiltonian formulation for the motion of an active spheroidal particle suspended in laminar straight duct flow
- Effects of Near-Field Hydrodynamic Interactions on Bacterial Dynamics Near a Solid Surface
- From Propulsion to Suction: Unraveling Thrust Reversal in Propellers at Intermediate Reynolds Numbers
- Reinforcement learning of a biflagellate model microswimmer
- Hydrodynamic surrogate models for bio-inspired micro-swimming robots
- Modal analysis and optimization of swimming active filaments
- The asymptotic coarse-graining formulation of slender-rods, bio-filaments and flagella
- Complete analytic solution to Brownian unicycle dynamics
- Planar Rotational Equilibria of Two Non-identical Microswimmers
- Algebraic correlations and anomalous fluctuations in ordered flocks of Janus particles fueled by an AC electric field
- Simple Flagellated Soft Robot for Locomotion near Air-Fluid Interface
- Effective squirmer models for self-phoretic chemically active spherical colloids
- Chiral propulsion by electromagnetic fields
- Simulating squirmers with volumetric solvers
- Swimming of microorganisms in quasi-2D membranes
- Analysis of aligning active local searchers orbiting around their common home position
- Irreversibility of stochastic state transitions in Langevin systems with odd elasticity
- Dynamic Force Measurements on Swimming Chlamydomonas Cells using Micropipette Force Sensors
- Symmetry-group-protected microfluidics for multiplexed stress-free manipulations
- On the solidity parameter in canopy flows
- An integral model based on slender body theory, with applications to curved rigid fibers
- Optimal Strokes of Low Reynolds Number Linked-Sphere Swimmers
- Bayesian uncertainty quantification for micro-swimmers with fully resolved hydrodynamics
- Dispersion Relations for Active Undulators in Overdamped Environments
- Inertial dynamics of an active Brownian particle
- Diffusion of Chiral Janus Particles in Convection Rolls
- Wall attraction and repulsion of hydrodynamically interacting particles
- A Swimming Rheometer: Self-propulsion of a freely-suspended swimmer enabled by viscoelastic normal stresses
- Mass-based separation of active Brownian particles in an asymmetric channel
- Surfing on chemical waves: a simple yet dynamically rich two-sphere responsive gel swimmer
- Control of Microparticles Through Hydrodynamic Interactions
- Effect of Convection Rolls in Motility-Induced Phase Separation of Active Janus Particles
- Generalized squirming motion of a sphere
- Dynamics of cruising swimming by a deformable sphere for two simple models
- Beyond the Markovian limit: Exact solutions for active motion in a power-law viscoelastic bath
- Optimal dynamics of a spherical squirmer in Eulerian description
- Experimental investigations on the nonequilibrium dynamics of pattern formation in fluid and granular systems
- Shear induced migration of microswimmers in pressure-driven channel flow
- A three-dimensional morphoelastic model for self-oscillations in polyelectrolyte hydrogel filaments
- Learning to traverse convective flows at moderate to high Rayleigh numbers
- On the DLM/FD methods for simulating neutrally buoyant swimmer motion in non-Newtonian shear thinning fluids
- Large-scale simulation of steady and time-dependent active suspensions with the force-coupling method
- Simulating cilia-driven mixing and transport in complex geometries
- Hamiltonian Active Particles in Incompressible Fluid Membranes
- Active Caustics
- Method of regularised stokeslets: Flow analysis and improvement of convergence
- Propulsion and controlled steering of magnetic nanohelices
- A swimming bacterium in a two-fluid model of a polymer solution
- Hydrodynamics of flagellated microswimmers near free-slip interfaces
- Paramecium swimming in capillary tube
- Undulatory swimming in suspensions and networks of flexible filaments
- Mixing by microorganisms in stratified fluids
- Diffusion of gravitactic chiral active Brownian particles in an asymmetric channel
- Enhanced locomotion, effective diffusion, and trapping of undulatory micro-swimmers in heterogeneous environments
- Dynamics of a simple model microswimmer in an anisotropic fluid: implications for alignment behavior and active transport in a nematic liquid crystal
- Density-Independent transient caging in the high-density phase of motility-induced phase separation
- Microswimming in viscoelastic fluids
- Three-dimensional vibrations of multilayered hollow spheres submerged in a complex fluid
- Travelling waves are hydrodynamically optimal for long-wavelength flagella
- Direct vs indirect hydrodynamic interactions during bundle formation of bacterial flagella
- Directional motion of forced polymer chains with hydrodynamic interaction
- Nonlinear dynamics and fluctuations in biological systems (Habilitation thesis)
- Enhanced Diffusion and Chemotaxis of Enzymes
- Base flow decomposition for complex moving objects in linear hydrodynamics: application to helix-shaped flagellated microswimmers
- Pairwise and collective behavior between model swimmers at intermediate Reynolds numbers