In order to understand the rheological and transport properties of a suspension of swimming micro-organisms, it is necessary to analyse the fluid-dynamical interaction of pairs of such swimming cells. In this paper, a swimming micro-organism is modelled as a squirming sphere with prescribed tangential surface velocity, referred to as a squirmer. The centre of mass of the sphere may be displaced from the geometric centre (bottom-heaviness). The effects of inertia and Brownian motion are neglected, because real micro-organisms swim at very low Reynolds numbers but are too large for Brownian effects to be important. The interaction of two squirmers is calculated analytically for the limits of small and large separations and is also calculated numerically using a boundary-element method. The analytical and the numerical results for the translational–rotational velocities and for the stresslet of two squirmers correspond very well. We sought to generate a database for an interacting pair of squirmers from which one can easily predict the motion of a collection of squirmers. The behaviour of two interacting squirmers is discussed phenomenologically, too. The results for the trajectories of two squirmers show that first the squirmers attract each other, then they change their orientation dramatically when they are in near contact and finally they separate from each other. The effect of bottom-heaviness is considerable. Restricting the trajectories to two dimensions is shown to give misleading results. Some movies of interacting squirmers are available with the online version of the paper.

The spherical alga Volvox swims by means of flagella on thousands of surface somatic cells. This geometry and its large size make it a model organism for studying the fluid dynamics of multicellularity. Remarkably, when two nearby Volvox colonies swim close to a solid surface, they attract one another and can form stable bound states in which they “waltz” or “minuet” around each other. A surface-mediated hydrodynamic attraction combined with lubrication forces between spinning, bottom-heavy Volvox explains the formation, stability, and dynamics of the bound states. These phenomena are suggested to underlie observed clustering of Volvox at surfaces.

We report various types of coherent structures in suspensions of spherical particles swimming in a monolayer. We solve the fluid dynamics precisely from far-field hydrodynamic interactions to lubrication between two near-contact surfaces. The simulation results clearly illustrate that coherent structures, such as aggregation, mesoscale spatiotemporal motion, and band formation, can be generated by purely hydrodynamic interactions.

The rheological properties of a cell suspension may play an important role in the flow field generated by populations of swimming micro-organisms (e.g. in bioconvection). In this paper, a swimming micro-organism is modelled as a squirming sphere with prescribed tangential surface velocity, in which the centre of mass of the sphere may be displaced from the geometric centre (bottom-heaviness). Effects of inertia and Brownian motion are neglected, because real micro-organisms swim at very low Reynolds numbers but are too large for Brownian effects to be important. The three-dimensional movement of 64 identical squirmers in a simple shear flow field, contained in a cube with periodic boundary conditions, is dynamically computed, for random initial positions and orientations. The computation utilizes a database of pairwise interactions that has been constructed by the boundary element method. The restriction to pairwise additivity of forces is expected to be justified if the suspension is semi-dilute. The results for non-bottom-heavy squirmers show that the squirming does not have a direct influence on the apparent viscosity. However, it does change the probability density in configuration space, and thereby causes a slight decrease in the apparent viscosity atO(c²), wherecis the volume fraction of spheres. In the case of bottom-heavy squirmers, on the other hand, the stresslet generated by the squirming motion directly contributes to the bulk stress atO(c), and the suspension shows strong non-Newtonian properties. When the background simple shear flow is directed vertically, the apparent viscosity of the semi-dilute suspension of bottom-heavy squirmers becomes smaller than that of inert spheres. When the shear flow is horizontal and varies with the vertical coordinate, on the other hand, the apparent viscosity becomes larger than that of inert spheres. In addition, significant normal stress differences appear for all relative orientations of gravity and the shear flow, in the case of bottom-heavy squirmers.

The diffusive behaviour of swimming micro-organisms should be clarified in order to obtain a better continuum model for cell suspensions. In this paper, a swimming micro-organism is modelled as a squirming sphere with prescribed tangential surface velocity, in which the centre of mass of the sphere may be displaced from the geometric centre (bottom-heaviness). Effects of inertia and Brownian motion are neglected, because real micro-organisms swim at very low Reynolds numbers but are too large for Brownian effects to be important. The three-dimensional movement of 64 or 27 identical squirmers in a fluid otherwise at rest, contained in a cube with periodic boundary conditions, is dynamically computed, for random initial positions and orientations. The computation utilizes a database of pairwise interactions that has been constructed by the boundary element method. In the case of (non-bottom-heavy) squirmers, both the translational and the orientational spreading of squirmers is correctly described as a diffusive process over a sufficiently long time scale, even though all the movements of the squirmers were deterministically calculated. Scaling of the results on the assumption that the squirmer trajectories are unbiased random walks is shown to capture some but not all of the main features of the results. In the case of (bottom-heavy) squirmers, the diffusive behaviour in squirmers' orientations can be described by a biased random walk model, but only when the effect of hydrodynamic interaction dominates that of the bottom-heaviness. The spreading of bottom-heavy squirmers in the horizontal directions show diffusive behaviour, and that in the vertical direction also does when the average upward velocity is subtracted. The rotational diffusivity in this case, at a volume fractionc=0.1, is shown to be at least as large as that previously measured in very dilute populations of swimming algal cells (Chlamydomonas nivalis).

A swimming micro-organism is modelled as a squirming sphere with prescribed tangential surface velocity and referred to as a squirmer. The centre of mass of the sphere may be displaced from the geometric centre, and the effects of inertia and Brownian motion are neglected. The well-known Stokesian dynamics method is modified in order to simulate squirmer motions in a concentrated suspension. The movement of 216 identical squirmers in a concentrated suspension without any imposed flow is simulated in a cubic domain with periodic boundary conditions, and the coherent structures within the suspension are investigated. The results show that (a) a weak aggregation of cells appears as a result of the hydrodynamic interaction between cells; (b) the cells generate collective motions by the hydrodynamic interaction between themselves; and (c) the range and duration of the collective motions depend on the volume fraction and the squirmers' stresslet strengths. These tendencies show good qualitative agreement with previous experiments.