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.
Escherichia coli is a motile bacterium that moves up a chemoattractant gradient by performing a biased random walk composed of alternating runs and tumbles. Previous models of run and tumble chemotaxis neglect one or more features of the motion, namely (a) a cell cannot directly detect a chemoattractant gradient but rather makes temporal comparisons of chemoattractant concentration, (b) rather than being entirely random, tumbles exhibit persistence of direction, meaning that the new direction after a tumble is more likely to be in the forward hemisphere, and (c) rotational Brownian motion makes it impossible for an E. coli cell to swim in a straight line during a run. This paper presents an analytic calculation of the chemotactic drift velocity taking account of (a), (b) and (c), for weak chemotaxis. The analytic results are verified by Monte Carlo simulation. The results reveal a synergy between temporal comparisons and persistence that enhances the drift velocity, while rotational Brownian motion reduces the drift velocity.
We calculate non-Brownian fluid particle diffusion in a semidilute suspension of swimming micro-organisms. Each micro-organism is modeled as a spherical squirmer, and their motions in an infinite suspension otherwise at rest are computed by the Stokesian-dynamics method. In calculating the fluid particle motions, we propose a numerical method based on a combination of the boundary element technique and Stokesian dynamics. We present details of the numerical method and examine its accuracy. The limitation of semidiluteness is required to ensure accuracy of the fluid particle velocity calculation. In the case of a suspension of non-bottom-heavy squirmers the spreading of fluid particles becomes diffusive in a shorter time than that of the squirmers, and the diffusivity of fluid particles is smaller than that of squirmers. It is confirmed that the probability density distribution of fluid particles also shows diffusive properties. The effect of tracer particle size is investigated by inserting some inert spheres of the same radius as the squirmers, instead of fluid particles, into the suspension. The diffusivity for inert spheres is not less than one tenth of that for fluid particles, even though the particle size is totally different. Scaling analysis indicates that the diffusivity of fluid particles and inert spheres becomes proportional to the volume fraction of squirmers in the semidilute regime provided that there is no more than a small recirculation region around a squirmer, which is confirmed numerically. In the case of a suspension of bottom-heavy squirmers, horizontal diffusivity decreases considerably even with small values of the bottom heaviness, which indicates the importance of bottom heaviness in the diffusion phenomena. We believe that these fundamental findings will enhance our understanding of the basic mechanics of a suspension of swimming micro-organisms.
Escherichia coli is a motile bacterium that moves up a chemoattractant gradient by performing a biased random walk composed of alternating runs and tumbles. This paper presents calculations of the chemotactic drift velocity v (d) (the mean velocity up the chemoattractant gradient) of an E. coli cell performing chemotaxis in a uniform, steady shear flow, with a weak chemoattractant gradient at right angles to the flow. Extending earlier models, a combined analytic and numerical approach is used to assess the effect of several complications, namely (i) a cell cannot detect a chemoattractant gradient directly but rather makes temporal comparisons of chemoattractant concentration, (ii) the tumbles exhibit persistence of direction, meaning that the swimming directions before and after a tumble are correlated, (iii) the cell suffers random re-orientations due to rotational Brownian motion, and (iv) the non-spherical shape of the cell affects the way that it is rotated by the shear flow. These complications influence the dependence of v(d) on the shear rate gamma. When they are all included, it is found that (a) shear disrupts chemotaxis and shear rates beyond gamma approximately 2 s(-1) render chemotaxis ineffective, (b) in terms of maximizing drift velocity, persistence of direction is advantageous in a quiescent fluid but disadvantageous in a shear flow, and (c) a more elongated body shape is advantageous in performing chemotaxis in a shear flow.
Previously published experimental work by other authors has shown that certain motile marine bacteria are able to track free-swimming algae by executing a zigzag path and steering toward the algae at each turn. Here, we propose that the apparent steering behaviour could be a hydrodynamic effect, whereby an algal cell's vorticity and strain-rate fields rotate a pursuing bacterial cell in the appropriate direction. Using simplified models for the bacterial and algal cells, we numerically compute the trajectory of a bacterial cell and demonstrate the plausibility of this hypothesis.
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