The Hydrodynamics of Active Systems

This is a series of four lectures presented at the 2015 Enrico Fermi summer school in Varenna. The aim of the lectures is to give an introduction to the hydrodynamics of active matter concentrating on low Reynolds number examples such as cells and molecular motors. Lecture 1 introduces the hydrodynamics of single active particles, covering the Stokes equation and the Scallop Theorem, and stressing the link between autonomous activity and the dipolar symmetry of the far flow field. In lecture 2 I discuss applications of this mathematics to the behaviour of microswimmers at surfaces and in external flows, and describe our current understanding of how swimmers stir the surrounding fluid. Lecture 3 concentrates on the collective behaviour of active particles, modelled as an active nematic. I write down the equations of motion and motivate the form of the active stress. The resulting hydrodynamic instability leads to a state termed active turbulence characterised by strong jets and vortices in the flow field and the continual creation and annihilation of pairs of topological defects. Lecture 4 compares simulations of active turbulence to experiments on suspensions of microtubules and molecular motors. I introduce lyotropic active nematics and discuss active anchoring at interfaces.Comment: Lecture Notes, 2015 Enrico Fermi Summer School on Soft Matter Self-Assembly, Vienn

Entrainment and scattering in microswimmer--colloid interactions

We use boundary element simulations to study the interaction of model microswimmers with a neutrally buoyant spherical particle. The ratio of the size of the particle to that of the swimmer is varied from $R^\mathrm{P} / R^\mathrm{S} \ll 1$, corresponding to swimmer--tracer scattering, to $R^\mathrm{P} / R^\mathrm{S} \gg 1$, approximately equivalent to the swimmer interacting with a fixed, flat surface. We find that details of the swimmer and particle trajectories vary for different swimmers. However, the overall characteristics of the scattering event fall into two regimes, depending on the relative magnitudes of the impact parameter, $\rho$, and the collision radius, $R^\mathrm{coll}=R^\mathrm{P} + R^\mathrm{S}$. The range of particle motion, defined as the maximum distance between two points on the trajectory, has only a weak dependence on the impact parameter when $\rho < R^\mathrm{coll}$ and decreases with the radius of the particle. In contrast, when $\rho>R^\mathrm{coll}$ the range decreases as a power law in $\rho$ and is insensitive to the size of the particle. We also demonstrate that large particles can cause swimmers to be deflected through large angles. In some instances, this swimmer deflection can lead to larger net displacements of the particle. Based on these results, we estimate the effective diffusivity of a particle in a dilute bath of swimmers and show that there is a non-monotonic dependence on particle radius. Similarly, we show that the effective diffusivity of a swimmer scattering in a suspension of particles varies non-monotonically with particle radius.Comment: 19 pages, 11 figures. Accepted in Physical Review Fluid

Fluid mixing by curved trajectories of microswimmers

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 $\lambda$, we obtain an exact expression for $D_{rr}$ for dipolar swimmers in three dimensions, hence explaining the surprising result that this is independent of $\lambda$. We compare $D_{rr}$ to the contribution to tracer diffusion from entrainment.Comment: 5 pages, 2 figure

Stirring by swimmers in confined microenvironments

We consider the tracer diffusion $D_{rr}$ that arises from the run-and-tumble motion of low Reynolds number swimmers, such as bacteria. In unbounded dilute suspensions, where the dipole swimmers move in uncorrelated runs of length $\lambda$, an exact solution showed that $D_{rr}$ is independent of $\lambda$. Here we verify this result in numerical simulations for a particular model swimmer, the spherical squirmer. We also note that in confined microenvironments, such as microscopic droplets, microfluidic devices and bacterial microzones in marine ecosystems, the size of the system can be comparable to $\lambda$. We show that this effect alone reduces the value of $D_{rr}$ in comparison to its bulk value, and predict a scaling form for its relative decrease.Comment: submitted to JSTA

Multi-scale statistics of turbulence motorized by active matter

A number of micro-scale biological flows are characterized by spatio-temporal chaos. These include dense suspensions of swimming bacteria, microtubule bundles driven by motor proteins, and dividing and migrating confluent layers of cells. A characteristic common to all of these systems is that they are laden with active matter, which transforms free energy in the fluid into kinetic energy. Because of collective effects, the active matter induces multi-scale flow motions that bear strong visual resemblance to turbulence. In this study, multi-scale statistical tools are employed to analyze direct numerical simulations (DNS) of periodic two- (2D) and three-dimensional (3D) active flows and compare them to classic turbulent flows. Statistical descriptions of the flows and their variations with activity levels are provided in physical and spectral spaces. A scale-dependent intermittency analysis is performed using wavelets. The results demonstrate fundamental differences between active and high-Reynolds number turbulence; for instance, the intermittency is smaller and less energetic in active flows, and the work of the active stress is spectrally exerted near the integral scales and dissipated mostly locally by viscosity, with convection playing a minor role in momentum transport across scales.Comment: Accepted in Journal of Fluid Mechanics (2017

CUDA simulations of active dumbbell suspensions

We describe and analyze CUDA simulations of hydrodynamic interactions in active dumbbell suspensions. GPU-based parallel computing enables us not only to study the time-resolved collective dynamics of up to a several hundred active dumbbell swimmers but also to test the accuracy of effective time-averaged models. Our numerical results suggest that the stroke-averaged model yields a relatively accurate description down to distances of only a few times the dumbbell's length. This is remarkable in view of the fact that the stroke-averaged model is based on a far-field expansion. Thus, our analysis confirms that stroke-averaged far-field equations of motion may provide a useful starting point for the derivation of hydrodynamic field equations.Comment: 16 pages, 4 figure