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Register a new userThe Bank Of Swimming Organisms at the Micron Scale (BOSO-Micro)
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Unicellular microscopic organisms living in aqueous environments outnumber all other creatures on Earth. A large proportion of them are able to self-propel in fluids with a vast diversity of swimming gaits and motility patterns. In this paper we present a biophysical survey of the available experimental data produced to date on the characteristics of motile behaviour in unicellular microswimmers. We assemble from the available literature empirical data on the motility of four broad categories of organisms: bacteria (and archaea), flagellated eukaryotes, spermatozoa and ciliates. Whenever possible, we gather the following biological, morphological, kinematic and dynamical parameters: species, geometry and size of the organisms, swimming speeds, actuation frequencies, actuation amplitudes, number of flagella and properties of the surrounding fluid. We then organise the data using the established fluid mechanics principles for propulsion at low Reynolds number. Specifically, we use theoretical biophysical models for the locomotion of cells within the same taxonomic groups of organisms as a means of rationalising the raw material we have assembled, while demonstrating the variability for organisms of different species within the same group. The material gathered in our work is an attempt to summarise the available experimental data in the field, providing a convenient and practical reference point for future studies.
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Many microorganisms and artificial microswimmers use helical appendages in order to generate locomotion. Though often rotated so as to produce thrust, some species of bacteria such Spiroplasma, Rhodobacter sphaeroides and Spirochetes induce movement by deforming a helical-shaped body. Recently, artificial devices have been created which also generate motion by deforming their helical body in a non-reciprocal way (Mourran et al., Adv. Mater., 29, 1604825, 2017). Inspired by these systems, we investigate the transport of a deforming helix within a viscous fluid. Specifically, we consider a swimmer that maintains a helical centreline and a single handedness while changing its helix radius, pitch and wavelength uniformly across the body. We first discuss how a deforming helix can create a non-reciprocal translational and rotational swimming stroke and identify its principle direction of motion. We then determine the leading-order physics for helices with small helix radius before considering the general behaviour for different configuration parameters and how these swimmers can be optimised. Finally, we explore how the presence of walls, gravity, and defects in the centreline allow the helical device to break symmetries, increase its speed, and generate transport in directions not available to helices in bulk fluids.
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.
In a variety of biological processes, eukaryotic cells use cilia to transport flow. Although cilia have a remarkably conserved internal molecular structure, experimental observations report very diverse kinematics. To address this diversity, we determine numerically the kinematics and energetics of the most efficient cilium. Specifically, we compute the time-periodic deformation of a wall-bound elastic filament leading to transport of a surrounding fluid at minimum energetic cost, where the cost is taken to be the positive work done by all internal molecular motors. The optimal kinematics are found to strongly depend on the cilium bending rigidity through a single dimensionless number, the Sperm number, and closely resemble the two-stroke ciliary beating pattern observed experimentally.
Peritrichous bacteria such as Escherichia coli swim in viscous fluids by forming a helical bundle of flagellar filaments. The filaments are spatially distributed around the cell body to which they are connected via a flexible hook. To understand how the swimming direction of the cell is determined, we theoretically investigate the elastohydrodynamic motility problem of a multi-flagellated bacterium. Specifically, we consider a spherical cell body with a number N of flagella which are initially symmetrically arranged in a plane in order to provide an equilibrium state. We analytically solve the linear stability problem and find that at most 6 modes can be unstable and that these correspond to the degrees of freedom for the rigid-body motion of the cell body. Although there exists a rotation-dominated mode that generates negligible locomotion, we show that for the typical morphological parameters of bacteria the most unstable mode results in linear swimming in one direction accompanied by rotation around the same axis, as observed experimentally.
In a classic paper, Edward Purcell analysed the dynamics of flagellated bacterial swimmers and derived a geometrical relationship which optimizes the propulsion efficiency. Experimental measurements for wild-type bacterial species E. coli have revealed that they closely satisfy this geometric optimality. However, the dependence of the flagellar motor speed on the load and more generally the role of the torque-speed characteristics of the flagellar motor is not considered in Purcells original analysis. Here we derive a tuned condition representing a match between the flagella geometry and the torque-speed characteristics of the flagellar motor to maximize the bacterial swimming speed for a given load. This condition is independent of the geometric optimality condition derived by Purcell and interestingly this condition is not satisfied by wild-type E. coli which swim 2-3 times slower than the maximum possible speed given the amount of available motor torque. Our analysis also reveals the existence of an anomalous propulsion regime, where the swim speed increases with increasing load (drag). Finally, we present experimental data which supports our analysis.
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