Hydrodynamic Irreversibility in Particle Suspensions with Non-Uniform Strain

A dynamical phase transition from reversible to irreversible behavior occurs when particle suspensions are subjected to uniform oscillatory shear, even in the Stokes flow limit. We consider a more general situation with non-uniform strain (e.g. oscillatory channel flow), which is observed to exhibit markedly different dynamics. Self-organization and shear-induced migration only partially explain the delayed, simultaneous onset of irreversibility across the channel. The onset of irreversibility is accompanied by long-range correlated particle motion. This motion leads to particle activity even at the channel center, where the strain is negligible, and prevents the system from evolving into a reversible state

Oscillatory Flows Induced by Microorganisms Swimming in Two-dimensions

We present the first time-resolved measurements of the oscillatory velocity field induced by swimming unicellular microorganisms. Confinement of the green alga C. reinhardtii in stabilized thin liquid films allows simultaneous tracking of cells and tracer particles. The measured velocity field reveals complex time-dependent flow structures, and scales inversely with distance. The instantaneous mechanical power generated by the cells is measured from the velocity fields and peaks at 15 fW. The dissipation per cycle is more than four times what steady swimming would require.Comment: 4 pages, 4 figure

Mixing by Swimming Algae

In this fluid dynamics video, we demonstrate the microscale mixing enhancement of passive tracer particles in suspensions of swimming microalgae, Chlamydomonas reinhardtii. These biflagellated, single-celled eukaryotes (10 micron diameter) swim with a "breaststroke" pulling motion of their flagella at speeds of about 100 microns/s and exhibit heterogeneous trajectory shapes. Fluorescent tracer particles (2 micron diameter) allowed us to quantify the enhanced mixing caused by the swimmers, which is relevant to suspension feeding and biogenic mixing. Without swimmers present, tracer particles diffuse slowly due solely to Brownian motion. As the swimmer concentration is increased, the probability density functions (PDFs) of tracer displacements develop strong exponential tails, and the Gaussian core broadens. High-speed imaging (500 Hz) of tracer-swimmer interactions demonstrates the importance of flagellar beating in creating oscillatory flows that exceed Brownian motion out to about 5 cell radii from the swimmers. Finally, we also show evidence of possible cooperative motion and synchronization between swimming algal cells.Comment: 1 page, APS-DFD 2009 Gallery of Fluid Motio

Direct measurement of the flow field around swimming microorganisms

Swimming microorganisms create flows that influence their mutual interactions and modify the rheology of their suspensions. While extensively studied theoretically, these flows have not been measured in detail around any freely-swimming microorganism. We report such measurements for the microphytes Volvox carteri and Chlamydomonas reinhardtii. The minute ~0.3% density excess of V. carteri over water leads to a strongly dominant Stokeslet contribution, with the widely-assumed stresslet flow only a correction to the subleading source dipole term. This implies that suspensions of V. carteri have features similar to suspensions of sedimenting particles. The flow in the region around C. reinhardtii where significant hydrodynamic interaction is likely to occur differs qualitatively from a "puller" stresslet, and can be described by a simple three-Stokeslet model.Comment: 4 pages, 4 figures, accepted for publication in PR

Nanoparticle image velocimetry at topologically structured surfaces

Nanoparticle image velocimetry ͑nano-PIV͒, based on total internal reflection fluorescent microscopy, is very useful to investigate fluid flows within ϳ100 nm from a surface; but so far it has only been applied to flow over smooth surfaces. Here we show that it can also be applied to flow over a topologically structured surface, provided that the surface structures can be carefully configured not to disrupt the evanescent-wave illumination. We apply nano-PIV to quantify the flow velocity distribution over a polydimethylsiloxane surface, with a periodic gratinglike structure ͑with 215 nm height and 2 m period͒ fabricated using our customized multilevel lithography method. The measured tracer displacement data are in good agreement with the computed theoretical values. These results demonstrate new possibilities to study the interactions between fluid flow and topologically structured surfaces

Periodic and Quasiperiodic Motion of an Elongated Microswimmer in Poiseuille Flow

We study the dynamics of a prolate spheroidal microswimmer in Poiseuille flow for different flow geometries. When moving between two parallel plates or in a cylindrical microchannel, the swimmer performs either periodic swinging or periodic tumbling motion. Although the trajectories of spherical and elongated swimmers are qualitatively similar, the swinging and tumbling frequency strongly depends on the aspect ratio of the swimmer. In channels with reduced symmetry the swimmers perform quasiperiodic motion which we demonstrate explicitely for swimming in a channel with elliptical cross section

Fluid dynamics and noise in bacterial cell-cell and cell-surface scattering

Bacterial processes ranging from gene expression to motility and biofilm formation are constantly challenged by internal and external noise. While the importance of stochastic fluctuations has been appreciated for chemotaxis, it is currently believed that deterministic long-range fluid dynamical effects govern cell-cell and cell-surface scattering - the elementary events that lead to swarming and collective swimming in active suspensions and to the formation of biofilms. Here, we report the first direct measurements of the bacterial flow field generated by individual swimming Escherichia coli both far from and near to a solid surface. These experiments allowed us to examine the relative importance of fluid dynamics and rotational diffusion for bacteria. For cell-cell interactions it is shown that thermal and intrinsic stochasticity drown the effects of long-range fluid dynamics, implying that physical interactions between bacteria are determined by steric collisions and near-field lubrication forces. This dominance of short-range forces closely links collective motion in bacterial suspensions to self-organization in driven granular systems, assemblages of biofilaments, and animal flocks. For the scattering of bacteria with surfaces, long-range fluid dynamical interactions are also shown to be negligible before collisions; however, once the bacterium swims along the surface within a few microns after an aligning collision, hydrodynamic effects can contribute to the experimentally observed, long residence times. As these results are based on purely mechanical properties, they apply to a wide range of microorganisms.Comment: 9 pages, 2 figures, http://www.pnas.org/content/108/27/1094

Cell morphology governs directional control in swimming bacteria

The ability to rapidly detect and track nutrient gradients is key to the ecological success of motile bacteria in aquatic systems. Consequently, bacteria have evolved a number of chemotactic strategies that consist of sequences of straight runs and reorientations. Theoretically, both phases are affected by fluid drag and Brownian motion, which are themselves governed by cell geometry. Here, we experimentally explore the effect of cell length on control of swimming direction. We subjected Escherichia coli to an antibiotic to obtain motile cells of different lengths, and characterized their swimming patterns in a homogeneous medium. As cells elongated, angles between runs became smaller, forcing a change from a run-and-tumble to a run-and-stop/reverse pattern. Our results show that changes in the motility pattern of microorganisms can be induced by simple morphological variation, and raise the possibility that changes in swimming pattern may be triggered by both morphological plasticity and selection on morphology

MicroMotility: State of the art, recent accomplishments and perspectives on the mathematical modeling of bio-motility at microscopic scales

Mathematical modeling and quantitative study of biological motility (in particular, of motility at microscopic scales) is producing new biophysical insight and is offering opportunities for new discoveries at the level of both fundamental science and technology. These range from the explanation of how complex behavior at the level of a single organism emerges from body architecture, to the understanding of collective phenomena in groups of organisms and tissues, and of how these forms of swarm intelligence can be controlled and harnessed in engineering applications, to the elucidation of processes of fundamental biological relevance at the cellular and sub-cellular level. In this paper, some of the most exciting new developments in the fields of locomotion of unicellular organisms, of soft adhesive locomotion across scales, of the study of pore translocation properties of knotted DNA, of the development of synthetic active solid sheets, of the mechanics of the unjamming transition in dense cell collectives, of the mechanics of cell sheet folding in volvocalean algae, and of the self-propulsion of topological defects in active matter are discussed. For each of these topics, we provide a brief state of the art, an example of recent achievements, and some directions for future research

Enhancement of biomixing by swimming algal cells in two-dimensional films

Fluid mixing in active suspensions of microorganisms is important to ecological phenomena and presents a fascinating stochastic process. We investigate the mixing produced by swimming unicellular algal cells (Chlamydomonas) in quasi-two-dimensional liquid films by simultaneously tracking the motion of the cells and that of microscopic passive tracer particles advected by the fluid. The reduced spatial dimension of the system leads to long-range flows and a surprisingly strong dependence of tracer transport on the concentration of swimmers, which is explored over a wide range. The mean square displacements are well described by a stochastic Langevin model, which is used to parameterize the mixing. The effective diffusion coefficient D grows rapidly with the swimmer concentration PHI as D ∼ PHI 3/2, as a result of the increasing frequency of tracer-swimmer interactions and the long-range hydrodynamic disturbances created by the swimmers. Conditional sampling of the tracer data based on the instantaneous swimmer position shows that the rapid growth of the diffusivity enhancement with concentration must be due to particle interactions with multiple swimmers simultaneously. Finally, the anomalous probability distributions of tracer displacements become Gaussian at high concentration, but manifest strong power-law tails at low concentration, while the tracer displacements always grow diffusively in time.National Science Foundation (U.S.) (Grant No. NSF DMR-0803153