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 micrometers and below. At this scale, inertia is unimportant and the Reynolds number is small. Our emphasis is on the simple physical picture and fundamental flow physics phenomena in this regime. We first give a brief overview of the mechanisms for swimming motility, and of 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, in particular early calculations of swimming kinematics with prescribed stroke and the application of resistive force theory and slender-body theory to flagellar locomotion. After examining 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. (Some figures in this article are in colour only in the electronic version) This article was invited by Christoph Schmidt.

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The hydrodynamics of swimming microorganisms

LAMMPS - a flexible simulation tool for particle-based materials modeling at the atomic, meso, and continuum scales

Locomotion and transport of microorganisms in fluids is an essential aspect of life. Search for food, orientation toward light, spreading of off-spring, and the formation of colonies are only possible due to locomotion. Swimming at the microscale occurs at low Reynolds numbers, where fluid friction and viscosity dominates over inertia. Here, evolution achieved propulsion mechanisms, which overcome and even exploit drag. Prominent propulsion mechanisms are rotating helical flagella, exploited by many bacteria, and snake-like or whip-like motion of eukaryotic flagella, utilized by sperm and algae. For artificial microswimmers, alternative concepts to convert chemical energy or heat into directed motion can be employed, which are potentially more efficient. The dynamics of microswimmers comprises many facets, which are all required to achieve locomotion. In this article, we review the physics of locomotion of biological and synthetic microswimmers, and the collective behavior of their assemblies. Starting from individual microswimmers, we describe the various propulsion mechanism of biological and synthetic systems and address the hydrodynamic aspects of swimming. This comprises synchronization and the concerted beating of flagella and cilia. In addition, the swimming behavior next to surfaces is examined. Finally, collective and cooperate phenomena of various types of isotropic and anisotropic swimmers with and without hydrodynamic interactions are discussed.

/pdf/physics-of-microswimmers-single-particle-motion-and-4vxoi085wf.pdf

Physics of microswimmers--single particle motion and collective behavior: a review.

This paper reviews applications of the lattice-Boltzmann method to simulations of particle-fluid suspensions. We first summarize the available simulation methods for colloidal suspensions together with some of the important applications of these methods, and then describe results from lattice-gas and lattice-Boltzmann simulations in more detail. The remainder of the paper is an update of previously published work,(69, 70) taking into account recent research by ourselves and other groups. We describe a lattice-Boltzmann model that can take proper account of density fluctuations in the fluid, which may be important in describing the short-time dynamics of colloidal particles. We then derive macro-dynamical equations for a collision operator with separate shear and bulk viscosities, via the usual multi-time-scale expansion. A careful examination of the second-order equations shows that inclusion of an external force, such as a pressure gradient, requires terms that depend on the eigenvalues of the collision operator. Alternatively, the momentum density must be redefined to include a contribution from the external force. Next, we summarize recent innovations and give a few numerical examples to illustrate critical issues. Finally, we derive the equations for a lattice-Boltzmann model that includes transverse and longitudinal fluctuations in momentum. The model leads to a discrete version of the Green–Kubo relations for the shear and bulk viscosity, which agree with the viscosities obtained from the macro-dynamical analysis. We believe that inclusion of longitudinal fluctuations will improve the equipartition of energy in lattice-Boltzmann simulations of colloidal suspensions.

Lattice-Boltzmann Simulations of Particle-Fluid Suspensions

/pdf/physics-of-microswimmers-single-particle-motion-and-2tjtc65h8s.pdf

Physics of Microswimmers - Single Particle Motion and Collective Behavior

A model problem is analysed to study the microscopic flow near the surface of two-dimensional porous media. In the idealized problem we consider axial flow through infinite and semi-infinite lattices of cylindrical inclusions. The effect of lattice geometry and inclusion shape on the permeability and surface flow are examined. Calculations show that the definition of a slip coefficient for a porous medium is meaningful only for extremely dilute systems. Brinkman's equation gives reasonable predictions for the rate of decay of the mean velocity for certain simple geometries, but fails for to predict the correct behaviour for media anisotropic in the plane normal to the flow direction.

Microscopic flow near the surface of two-dimensional porous media. Part 1. Axial flow

Numerical results are presented for planar sinusoidal waves (amplitude α, wave-number k). The average swimming speed and power consumption are computed for a wide range of the parameters. The optimal sine wave for minimizing power consumption is found to be a single wave with amplitude αk ≈ 1. The power consumption is found to be relatively insensitive to changes in the flagellar radius. The optimal flagellar length is found to be in the range L/A = 20–40. The instantaneous force distribution and flow field for a typical organism are presented. The trajectory of the organism through one cycle shows that a wave of constant amplitude may have the appearance of increasing amplitude owing to the yawing motion of the organism.The results are compared with those obtained using resistance coefficients. For organisms with small cell bodies (A/L = 0.05), the average swimming speed predicted by Gray-Hancock coefficients is accurate to within 10%. For large cell bodies (A/L = 0.2), the error in swimming speed is approximately 20%. The relative error in the predicted power consumption is 25–50%. For the coefficients suggested by Lighthill, the power is consistently underestimated. The Gray-Hancock coefficients underestimate the power for small cell bodies and overestimate it for large cell bodies.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/S0022112079002482

A hydrodynamic analysis of flagellar propulsion

A model problem is analysed to study the microscopic flow near the surface of porous media. In the idealized system, we consider two-dimensional media consisting of infinite and semi-infinite periodic lattices of cylindrical inclusions. In Part 1, results for axial flow were presented. In this work results for transverse flow are presented and discussed in the context of macroscopic approaches such as slip coefficients and Brinkman's equation.

Microscopic flow near the surface of two-dimensional porous media. Part 2. Transverse flow

A method is described for solving the integral equations governing Stokes flow in arbitrary two-dimensional domains. It is demonstrated that the boundary-integral method provides an accurate, efficient and easy-to-implement strategy for the solution of Stokes-flow problems. Calculations are presented for simple shear flow in a variety of geometries including cylindrical and rectangular, ridges and cavities. A full description of the flow field is presented including streamline patterns, velocity profiles and shear-stress distributions along the solid surfaces. The results are discussed with special relevance to convective transport processes in low-Reynolds-number flows.

Stokes flow in arbitrary two-dimensional domains: shear flow over ridges and cavities

The swimming of a micro-organism by the propagation of helical waves on a long slender flagellum is analysed. The model developed by Higdon (1979) is used to study the motion of an organism with a spherical cell body (radius A) propelled by a cylindrical flagellum (radius a, length L).The average swimming speed and power consumption are calculated for helical waves (amplitude α, wavenumber k). A wide range of parameter values is considered to determine the optimal swimming motion. The optimal helical wave has ak ≈ 1, corresponding to a pitch angle of 45°. The optimum number of waves on the flagellum increases as the flagellar length L/A increases, such that the optimum wavelength decreases as L/A increases. The efficiency is relatively insensitive to the flagellar radius a/A. The optimum flagellar length is L/A ≈ 10.The results are compared to calculations using two different forms of resistance coefficients. Gray-Hancock coefficients overestimate the swimming speed by approximately 20% and underestimate the power consumption by 50%. The coefficients suggested by Lighthill (1976) overestimate the swimming speed for large cell bodies (L/A 15) by 10%. The Lighthill coefficients underestimate the power consumption up to 50% for L/A 10. Overall, the Lighthill coefficients are superior to the Gray-Hancock coefficients in modelling swimming by helical waves.

J. J. L. Higdon

Papers

Microscopic flow near the surface of two-dimensional porous media. Part 1. Axial flow

A hydrodynamic analysis of flagellar propulsion

Microscopic flow near the surface of two-dimensional porous media. Part 2. Transverse flow

Stokes flow in arbitrary two-dimensional domains: shear flow over ridges and cavities

The hydrodynamics of flagellar propulsion: helical waves