No Arabic abstract
Confinement and wall effects are known to affect the kinematics and propulsive characteristics of swimming microorganisms. When a solid body is dragged through a viscous fluid at constant velocity, the presence of a wall increases fluid drag, and thus the net force required to maintain speed has to increase. In contrast, recent optical trapping experiments have revealed that the propulsive force generated by human spermatozoa is decreased by the presence of boundaries. Here, we use a series of simple models to analytically elucidate the propulsive effects of a solid boundary on passively actuated filaments and model flagella. For passive flexible filaments actuated periodically at one end, the presence of the wall is shown to increase the propulsive forces generated by the filaments in the case of displacement-driven actuation, while it decreases the force in the case of force-driven actuation. In the case of active filaments as models for eukaryotic flagella, we demonstrate that the manner in which a solid wall affects propulsion cannot be known a priori, but is instead a nontrivial function of the flagellum frequency, wavelength, its material characteristics, the manner in which the molecular motors self-organize to produce oscillations (prescribed activity model or self-organized axonemal beating model), and the boundary conditions applied experimentally to the tethered flagellum. In particular, we show that in some cases, the increase in fluid friction induced by the wall can lead to a change in the waveform expressed by the flagella which results in a decrease in their propulsive force.
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.
We discuss the flow field and propulsion velocity of active droplets, which are driven by body forces residing on a rigid gel. The latter is modelled as a porous medium which gives rise to permeation forces. In the simplest model, the Brinkman equation, the porous medium is characterised by a single length scale $ell$ --the square root of the permeability. We compute the flow fields inside and outside of the droplet as well as the energy dissipation as a function of $ell$. We furthermore show that there are optimal gel fractions, giving rise to maximal linear and rotational velocities. In the limit $elltoinfty$, corresponding to a very dilute gel, we recover Stokes flow. The opposite limit, $ellto 0$, corresponding to a space filling gel, is singular and not equivalent to Darcys equation, which cannot account for self-propulsion.
The near-surface swimming patterns of bacteria are strongly determined by the hydrodynamic interactions between bacteria and the surface, which trap bacteria in smooth circular trajectories that lead to inefficient surface exploration. Here, we show by combining experiments and a data-driven mathematical model that surface exploration of enterohemorrhagic Escherichia coli (EHEC) -- a pathogenic strain of E. coli causing serious illnesses such as bloody diarrhea -- results from a complex interplay between motility and transient surface adhesion events. These events allow EHEC to break the smooth circular trajectories and regulate their transport properties by the use stop-adhesion events that lead to a characteristic intermittent motion on surfaces. We find that the experimentally measured frequency of stop-adhesion events in EHEC is located at the value predicted by the developed mathematical model that maximizes bacterial surface diffusivity. We indicate that these results and the developed model apply to other bacterial strains on different surfaces, which suggests that swimming bacteria use transient adhesion to regulate surface motion.
In a multitude of lifes processes, cilia and flagella are found indispensable. Recently, the biflagellated chlorophyte alga Chlamydomonas has become a model organism for the study of ciliary coordination and synchronization. Here, we use high-speed imaging of single pipette-held cells to quantify the rich dynamics exhibited by their flagella. Underlying this variability in behaviour, are biological dissimilarities between the two flagella - termed cis and trans, with respect to a unique eyespot. With emphasis on the wildtype, we use digital tracking with sub-beat-cycle resolution to obtain limit cycles and phases for self-sustained flagellar oscillations. Characterizing the phase-synchrony of a coupled pair, we find that during the canonical swimming breaststroke the cis flagellum is consistently phase-lagged relative to, whilst remaining robustly phase-locked with, the trans flagellum. Transient loss of synchrony, or phase-slippage, may be triggered stochastically, in which the trans flagellum transitions to a second mode of beating with attenuated beat-envelope and increased frequency. Further, exploiting this algas ability for flagellar regeneration, we mechanically induced removal of one or the other flagellum of the same cell to reveal a striking disparity between the beating of the cis vs trans flagellum, in isolation. This raises further questions regarding the synchronization mechanism of Chlamydomonas.
Sheep are gregarious animals, and they often aggregate into dense, cohesive flocks, especially under stress. In this paper, we use image processing tools to analyze a publicly available aerial video showing a dense sheep flock moving under the stimulus of a shepherding dog. Inspired by the fluidity of the motion, we implement a hydrodynamics approach, extracting velocity fields, and measuring their propagation and correlations in space and time. We find that while the flock overall is stationary, significant dynamics happens at the edges, notably in the form of fluctuations propagating like waves, and large-scale correlations spanning the entire flock. These observations highlight the importance of incorporating interfacial dynamics, for instance in the form of line tension, when using a hydrodynamics framework to model the dynamics of dense, non-polarized swarms.