Microscopic lifeforms rarely locomote in an unconfined liquid. Solid boundaries representing biological membranes, other swimmers, or filaments much larger than the swimmer can represent elements of confinement. Due to the long range of hydrodynamic forces at this scale, the boundaries often have a dominant impact on the physics of locomotion. Here we extend previous work on locomotion of a swimmer with a prescribed stroke in confined isotropic fluids to anisotropic fluids, using the model of a nematic liquid crystal. The competition between elasticity, hydrodynamics, and anchoring conditions leads to a complex locomotion problem with unique transport properties. We examine this problem analytically and numerically for a model swimmer near a bounding wall which can itself also be elastic. For strong planar anchoring at a rigid wall, we find that the swimming speed goes to the isotropic Newtonian limit as the swimmer gets close to the wall, although the power required to maintain the swimmer’s speed depends on liquid crystal properties. We also report new findings on the swimming speed due to large-amplitude waveforms in unbounded liquid crystals.
Madison S. Krieger, Saverio E. Spagnolie and Thomas R. Powers, “Swimming in a confined liquid crystal”, arXiv, PDF
In evolutionary processes, population structure has a substantial effect on natural selection. Here, we analyze how motion of individuals affects constant selection in structured populations. Motion is relevant because it leads to changes in the distribution of types as mutations march toward fixation or extinction. We describe motion as the swapping of individuals on graphs, and also more generally as the shuffling of individuals between reproductive updates. Beginning with a one-dimensional graph, the cycle, we prove that motion suppresses natural selection for death-birth updating or for any process that combines birth-death and death-birth updating. If the rule is purely birth-death updating, no change in fixation probability appears in the presence of motion. We further investigate how motion affects evolution on the square lattice and on weighted graphs. In the latter case, we find that motion can be either an amplifier or a suppressor of natural selection. In some cases, whether it is one or the other can be a function of the relative reproductive rate, indicating that motion is a subtle and complex attribute of evolving populations.
Madison S. Krieger, A. McAvoy and M. A. Nowak, “Effects of motion in structured populations”, Arxiv, PDF
Microorganisms often encounter anisotropy, for example in mucus and biofilms. We study how anisotropy and elasticity of the ambient fluid affects the speed of a swimming microorganism with a prescribed stroke. Motivated by recent experiments on swimming bacteria in anisotropic environments, we extend a classical model for swimming microorganisms, the Taylor swimming sheet, actuated by small-amplitude traveling waves in a three-dimensional nematic liquid crystal without twist. We calculate the swimming speed and entrained volumetric flux as a function of the swimmer’s stroke properties as well as the elastic and rheological properties of the liquid crystal. These results are then compared to previous results on an analogous swimmer in a hexatic liquid crystal, indicating large differences in the cases of small Ericksen number and in a nematic fluid when the tumbling parameter is near the transition to a shear-aligning nematic. We also propose a novel method of swimming in a nematic fluid by passing a traveling wave of director oscillation along a rigid wall.
Madison S. Krieger, Saverio E. Spagnolie and Thomas R. Powers, “Microscale locomotion in a nematic liquid crystal”, Soft Matter, 2015, 11, 9115 – 9125, arXiv, PDF
When a microorganism begins swimming from rest in a Newtonian fluid such as water, it rapidly attains its steady-state swimming speed since changes in the velocity field spread quickly when the Reynolds number is small. However, swimming microorganisms are commonly found or studied in complex fluids. Because these fluids have long relaxation times, the time to attain the steady-state swimming speed can also be long. In this article we study the swimming startup problem in the simplest liquid crystalline fluid: a two-dimensional hexatic liquid crystal film. We study the dependence of startup time on anchoring strength and Ericksen number, which is the ratio of viscous to elastic stresses. For strong anchoring, the fluid flow starts up immediately but the liquid crystal field and swimming velocity attain their sinusoidal steady-state values after a time proportional to the relaxation time of the liquid crystal. When the Ericksen number is high, the behavior is the same as in the strong anchoring case for any anchoring strength. We also find that the startup time increases with the ratio of the rotational viscosity to the shear viscosity, and then ultimately saturates once the rotational viscosity is much greater than the shear viscosity.
Madison S. Krieger, Marcelo A. Dias and Thomas R. Powers, “Transient swimming in a hexatic liquid crystal”, Eur. Phys. J. E. 38, 94, arXiv, PDF
The swimming behavior of bacteria and other microorganisms is sensitive to the physical properties of the fluid in which they swim. Mucus, biofilms, and artificial liquid-crystalline solutions are all examples of fluids with some degree of anisotropy that are also commonly encountered by bacteria. In this article, we study how liquid-crystalline order affects the swimming behavior of a model swimmer. The swimmer is a one-dimensional version of G. I. Taylor’s swimming sheet: an infinite line undulating with small-amplitude transverse or longitudinal traveling waves. The fluid is a two-dimensional hexatic liquid-crystalline film. We calculate the power dissipated, swimming speed, and flux of fluid entrained as a function of the swimmer’s waveform as well as properties of the hexatic film, such as the rotational and shear viscosity, the Frank elastic constant, and the anchoring strength. The departure from isotropic behavior is greatest for large rotational viscosity and weak anchoring boundary conditions on the orientational order at the swimmer surface. We even find that if the rotational viscosity is large enough, the transverse-wave swimmer moves in the opposite direction relative to a swimmer in an isotropic fluid.
Madison S. Krieger, Saverio E. Spagnolie and Thomas R. Powers, “Locomotion and transport in a hexatic liquid crystal”, Phys. Rev. E 90, 052503, arXiv, PDF
Recent experiments have shown that many species of microorganisms leave a solid surface at a fixed angle determined by steric interactions and near-field hydrodynamics. This angle is completely independent of the incoming angle. For several collisions in a closed body this determines a unique type of billiard system, an aspecular billiard in which the outgoing angle is fixed for all collisions. We analyze such a system using numerical simulation of this billiard for varying tables and outgoing angles, and also utilize the theory of one-dimensional maps and wavefront dynamics. When applicable we cite results from and compare our system to similar billiard systems in the literature. We focus on examples from three broad classes: the ellipse, the Bunimovich billiards, and the Sinai billiards. The effect of a noisy outgoing angle is also discussed.
Madison S. Krieger, “Microorganism billiards in closed plane curves”, arXiv, PDF