Some microorganisms, such as spermatozoa, synchronize their flagella when swimming in close proximity. Using a simplified model (two infinite, parallel, two-dimensional waving sheets), we show that phase-locking arises from hydrodynamics forces alone, and has its origin in the front-back asymmetry of the geometry of their flagellar waveform. The time-evolution of the phase difference between co-swimming cells depends only on the nature of this geometrical asymmetry, and microorganisms can phase-lock into conformations which minimize or maximize energy dissipation.
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.
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 …
One approach to quantifying biological diversity consists of characterizing the statistical distribution of specific properties of a taxonomic group or habitat. Microorganisms living in fluid environments, and for whom motility is key, exploit propulsion resulting from a rich variety of shapes, forms, and swimming strategies. Here, we explore the variability of swimming speed for unicellular eukaryotes based on published data. The data naturally partitions into that from flagellates (with a small number of flagella) and from ciliates (with tens or more). Despite the morphological and size diff…
It has been shown that a nanoliter chamber separated by a wall of asymmetric obstacles can lead to an inhomogeneous distribution of self-propelled microorganisms. Although it is well established that this rectification effect arises from the interaction between the swimmers and the non-centrosymmetric pillars, here we demonstrate numerically that its efficiency is strongly dependent on the detailed dynamics of the individual microorganism. In particular, for the case of run-and-tumble dynamics, the distribution of run lengths, the rotational diffusion and the partial preservation of run orient…
Many microorganisms live and evolve in complex fluids. Examples include mammalian spermatozoa in cervical mucus, worms (e.g., \textit{C. elegans}) in wet soil, and bacteria (e.g., \textit{H. pylori}) in our stomach lining. Due to the presence of (bio)polymers and/or solids, such fluids often display nonlinear response to (shear) stresses including viscoelasticity and shear-rate dependent viscosity. The successful interaction between these microorganisms and their fluid environment is critical to the function of many biological processes including human reproduction, ecosystem dynamics, and the…
The dynamic interactions between pairs of swimming microorganisms underpin the collective behaviour of larger suspensions, but accurately calculating pairwise collisions has typically required the use of numerical simulations in which hydrodynamic interactions are fully resolved. In this paper, we utilise analytical expressions for forces and torques acting on two closely separated spherical squirmers -- accurate to second order in the ratio of cell-cell spacing to squirmer radius -- in order to calculate their scattering dynamics. Attention is limited to squirmers whose orientation vectors li…
We present a fast, high-throughput method for characterizing the motility of microorganisms in 3D based on standard imaging microscopy. Instead of tracking individual cells, we analyse the spatio-temporal fluctuations of the intensity in the sample from time-lapse images and obtain the intermediate scattering function (ISF) of the system. We demonstrate our method on two different types of microorganisms: bacteria, both smooth swimming (run only) and wild type (run and tumble) Escherichia coli, and the bi-flagellate alga Chlamydomonas reinhardtii. We validate the methodology using computer sim…
Interactions between microorganisms and their complex flowing environments are essential in many biological systems. We develop a model for microswimmer dynamics in non-Newtonian Poiseuille flows. We predict that swimmers in shear-thickening (-thinning) fluids migrate upstream more (less) quickly than in Newtonian fluids and demonstrate that viscoelastic normal stress differences reorient swimmers causing them to migrate upstream at the centreline, in contrast to well-known boundary accumulation in quiescent Newtonian fluids. Based on these observations, we suggest a sorting mechanism to selec…
Flagella and cilia are cellular appendages that inherit essential functions of microbial life including sensing and navigating the environment. In order to propel a swimming microorganism they displace the surrounding fluid by means of periodic motions, while precisely-timed modulations of their beating patterns enable the cell to steer towards or away from specific locations. Characterizing the dynamic forces, however, is challenging and typically relies on indirect experimental approaches. Here, we present direct in vivo measurements of the dynamic forces of motile Chlamydomonas reinhardtii …
Many microorganisms swim through gels and non-Newtonian fluids in their natural environments. In this paper, we focus on microorganisms which use flagella for propulsion. We address how swimming velocities are affected in nonlinearly viscoelastic fluids by examining the problem of an infinitely long cylinder with arbitrary beating motion in the Oldroyd-B fluid. We solve for the swimming velocity in the limit in which deflections of the cylinder from its straight configuration are small relative to the radius of the cylinder and the wavelength of the deflections; furthermore, the radius of the …
The incessant activity of swimming microorganisms has a direct physical effect on surrounding microscopic objects, leading to enhanced diffusion far beyond the level of Brownian motion with possible influences on the spatial distribution of non-motile planktonic species and particulate drifters. Here we study in detail the effect of eukaryotic flagellates, represented by the green microalga Chlamydomonas reinhardtii, on microparticles. Macro- and micro-scopic experiments reveal that microorganism-colloid interactions are dominated by rare close encounters leading to large displacements through…
Seeing new depths: Three-dimensional flow of a free-swimming alga
University of Minnesota（明尼苏达大学）；National Institute for Mathematical Sciences（韩国数学科学研究所）；Chung-Ang University（中央大学）；University at Buffalo（布法罗纽约州立大学）；University of Cincinnati（辛辛那提大学）
Many microorganisms swim through gels, materials with nonzero zero-frequency elastic shear modulus, such as mucus. Biological gels are typically heterogeneous, containing both a structural scaffold (network) and a fluid solvent. We analyze the swimming of an infinite sheet undergoing transverse traveling wave deformations in the "two-fluid" model of a gel, which treats the network and solvent as two coupled elastic and viscous continuum phases. We show that geometric nonlinearities must be incorporated to obtain physically meaningful results. We identify a transition between regimes where the …
We use the boundary element method to study the low-Reynolds number locomotion of a spherical model microorganism in a circular tube. The swimmer propels itself by tangen- tial or normal surface motion in a tube whose radius is on the order of the swimmer size. Hydrodynamic interactions with the tube walls significantly affect the average swimming speed and power consumption of the model microorganism. In the case of swimming parallel to the tube axis, the locomotion speed is always reduced (resp. increased) for swimmers with tangential (resp. normal) deformation. In all cases, the rate of wor…
The motility of microorganisms is influenced greatly by their hydrodynamic interactions with the fluidic environment they inhabit. We show by direct experimental observation of the bi-flagellated alga Chlamydomonas reinhardtii that fluid elasticity and viscosity strongly influence the beating pattern - the gait - and thereby control the propulsion speed. The beating frequency and the wave speed characterizing the cyclical bending are both enhanced by fluid elasticity. Despite these enhancements, the net swimming speed of the alga is hindered for fluids that are sufficiently elastic. The origin…
Many motile microorganisms adjust their swimming motion relative to the gravitational field and thus counteract sedimentation to the ground. This gravitactic behavior is often the result of an inhomogeneous mass distribution which aligns the microorganism similar to a buoy. However, it has been suggested that gravitaxis can also result from a geometric fore-rear asymmetry, typical for many self-propelling organisms. Despite several attempts, no conclusive evidence for such an asymmetry-induced gravitactic motion exists. Here, we study the motion of asymmetric self-propelled colloidal particles…
Efficient uptake of nutrients from the environment is an important component in the fitness of all microorganisms, and its dependence on size may reveal clues to the origins of evolutionary transitions to multicellularity. Because potential benefits in uptake rates must be viewed in the context of other costs and benefits of size, such as varying predation rates and the increased metabolic costs associated with larger and more complex body plans, the uptake rate itself is not necessarily that which is optimized by evolution. Uptake rates can be strongly dependent on local organism geometry and…
Marine microorganisms often reach high swimming speeds, either to take advantage of evanescent nutrient patches or to beat Brownian forces. Since this implies that a sizable part of their energetic budget must be allocated to motion, it is reasonable to assume that some fast-swimming microorganisms may increase their nutrient intake by increasing their speed v. We formulate a model to investigate this hypothesis and its consequences, finding the steady state solutions and analyzing their stability. Surprisingly, we find that even modest increases in nutrient absorption may lead to a significan…
Spermatozoa flagella are known to synchronize when swimming in close proximity. We use a model consisting of two-dimensional sheets propagating transverse waves of displacement to demonstrate that fluid forces lead to such synchronization passively. Using two distinct asymptotic descriptions (small amplitude and long wavelength), we derive the synchronizing dynamics analytically for ar- bitrarily shaped waveforms in Newtonian fluids, and show that phase locking will always occur for sufficiently asymmetric shapes. We characterize the effect of the geometry of the waveforms and the separation b…
Microorganisms can preferentially orient and move along gradients of a chemo-attractant (i.e., chemotax) while colonies of many microorganisms can collectively undergo complex dynamics in response to chemo-attractants that they themselves produce. For colonies or groups of micro-swimmers we investigate how an "auto-chemotactic" response that should lead to swimmer aggregation is affected by the non-trivial fluid flows that are generated by collective swimming. For this, we consider chemotaxis models based upon a hydrodynamic theory of motile suspensions that are fully coupled to chemo-attracta…
Max Planck Institute for the Physics of Complex Systems（马克斯·普朗克复杂系统物理研究所）；Brandeis University（布兰迪斯大学）；Center for Systems Biology Dresden（德累斯顿系统生物学中心）；TU Dresden（德累斯顿工业大学）
Geometric confinements are frequently encountered in soft matter systems and in particular significantly alter the dynamics of swimming microorganisms in viscous media. Surface-related effects on the motility of microswimmers can lead to important consequences in a large number of biological systems, such as biofilm formation, bacterial adhesion and microbial activity. On the basis of low-Reynolds-number hydrodynamics, we explore the state diagram of a three-sphere microswimmer under channel confinement in a slit geometry and fully characterize the swimming behavior and trajectories for neutra…
It has been known for some time that some microorganisms can swim faster in high-viscosity gel-forming polymer solutions. These gel-like media come to mimic highly viscous heterogeneous environment that these microorganisms encounter in-vivo. The qualitative explanation of this phenomena first offered by Berg and Turner [Nature (London) 278, 349 (1979)], suggests that propulsion enhancement is a result of flagellum pushing on quasi-rigid loose polymer network formed in some polymer solutions. Inspired by these observations, inertia-less propulsion in a heterogeneous viscous medium composed of …
Hydrodynamic interactions (HIs) are important in biophysics research because they influence both the collective and the individual behaviour of microorganisms and self-propelled particles. For instance, HIs at the micro-swimmer level determine the attraction or repulsion between individuals, and hence their collective behaviour. Meanwhile, HIs between swimming appendages (e.g. cilia and flagella) influence the emergence of swimming gaits, synchronised bundles and metachronal waves. In this study, we address the issue of HIs between slender filaments separated by a distance larger than their co…
