Micro-swimmer locomotion in heterogeneous media is increasingly relevant in biological physics due to the prevalence of microorganisms in complex environments. A model for such porous media is the Brinkman fluid which accounts for a sparse matrix of stationary obstacles via a linear resistance term in the momentum equation. We investigate two models for the locomotion and the flow field generated by a swimmer in such a medium. First, we analyze a dumbbell swimmer composed of two spring-connected spheres and driven by a flagellar force and derive its exact swimming velocity as a function of the Brinkman medium resistance, showing that the swimmer monotonically slows down as the medium drag monotonically increases. In the limit of no resistance the model reduces to the classical Stokes dipole swimmer, while finite resistance introduces hydrodynamic screening that attenuates long-range interactions. Additionally, we derive an analytical expression for the far-field flow generated by a Brinkmanlet force-dipole, which can be used for propulsive point-dipole swimmer models. Remarkably, this approximation reproduces the dumbbell swimmer's flow field in the far-field regime with high accuracy. These results provide new analytical tools for understanding locomotion in complex fluids and offer foundational insights for future studies on collective behavior in active and passive suspensions within porous or structured environments.
To achieve propulsion at low Reynolds number, a swimmer must deform in a way that is not invariant under time-reversal symmetry; this result is known as the scallop theorem. We show here that there is no many-scallop theorem. We demonstrate that two active particles undergoing reciprocal deformations can swim collectively; moreover, polar particles also experience effective long-range interactions. These results are derived for a minimal dimers model, and generalized to more complex geometries on the basis of symmetry and scaling arguments. We explain how such cooperative locomotion can be rea…
The bacterium Helicobacter pylori causes ulcers in the stomach of humans by invading mucus layers protecting epithelial cells. It does so by chemically changing the rheological properties of the mucus from a high-viscosity gel to a low-viscosity solution in which it may self-propel. We develop a two-fluid model for this process of swimming under self-generated confinement. We solve exactly for the flow and the locomotion speed of a spherical swimmer located in a spherically symmetric system of two Newtonian fluids whose boundary moves with the swimmer. We also treat separately the special case…
Inspired by recent experiments using synthetic microswimmers to manipulate droplets, we investigate the low-Reynolds-number locomotion of a model swimmer (a spherical squirmer) encapsulated inside a droplet of comparable size in another viscous fluid. Meditated solely by hydrodynamic interactions, the encaged swimmer is seen to be able to propel the droplet, and in some situations both remain in a stable co-swimming state. The problem is tackled using both an exact analytical theory and a numerical implementation based on boundary element method, with a particular focus on the kinematics of th…
Anisotropic viscous drag is usually believed to be a requirement for the low Reynolds number locomotion of slender bodies such as flagella and cilia. Here we show that locomotion under isotropic drag is possible for extensible slender bodies. After general considerations, a two-ring swimmer and a model dinoflagellate flagellum are studied analytically to illustrate how extensibility can be exploited for self-propulsion without drag anisotropy. This new degree of freedom could be useful for some complex swimmer geometries and locomotion in complex fluid environments where drag anisotropy is wea…
Micro-swimmers can serve as cargo carriers that move deep inside complex flow networks. When a school collectively entrains the surrounding fluid, their transport capacity can be enhanced. This effect is quantified with good agreement between experiments with self-propelled droplets and a confined Brinkman squirmer model. The volume of liquid entrained can be much larger than the droplet itself, amplifying the effective cargo capacity over an order of magnitude, even for dilute schools. Hence, biological and engineered swimmers can efficiently transport materials into confined environments.
We conduct experiments with flexible swimmers to address the impact of fluid viscoelasticity on their locomotion. The swimmers are composed of a magnetic head actuated in rotation by a frequency-controlled magnetic field and a flexible tail whose deformation leads to forward propulsion. We consider both viscous Newtonian and glucose-based Boger fluids with similar viscosities. We find that the elasticity of the fluid systematically enhances the locomotion speed of the swimmer, and that this enhancement increases with Deborah number. Using Particle Image Velocimetry to visualize the flow field,…
Swimming cells often have to self-propel through fluids displaying non-Newtonian rheology. While past theoretical work seems to indicate that stresses arising from complex fluids should systematically hinder low-Reynolds number locomotion, experimental observations suggest that locomotion enhancement is possible. In this paper we propose a physical mechanism for locomotion enhancement of microscopic swimmers in a complex fluid. It is based on the fact that micro-structured fluids will generically phase-separate near surfaces, leading to the presence of low-viscosity layers which promote slip a…
Swimming cells and microorganisms must often move though complex fluids that contain an immersed microstructure such as polymer molecules, or filaments. In many important biological processes, such as mammalian reproduction and bacterial infection, the size of the immersed microstructure is comparable to that of the swimming cells. This leads to discrete swimmer-microstructure interactions that alter the swimmer's path and speed. In this paper, we use a combination of detailed simulation and data-driven stochastic models to examine the motion of a planar undulatory swimmer in an environment of…
In the limit of zero Reynolds number (Re), swimmers propel themselves exploiting a series of non-reciprocal body motions. For an artificial swimmer, a proper selection of the power source is required to drive its motion, in cooperation with its geometric and mechanical properties. Although various external fields (magnetic, acoustic, optical, etc.) have been introduced, electric fields are rarely utilized to actuate such swimmers experimentally in unbounded space. Here we use uniform and static electric fields to demonstrate locomotion of a bi-flagellated sphere at low Re via Quincke rotation.…
In several biologically relevant situations, cell locomotion occurs in polymeric fluids with Weissenberg {number} larger than one. Here we present results of three-dimensional numerical simulations for the steady locomotion of a self-propelled body in a model polymeric (Giesekus) fluid at low Reynolds number. Locomotion is driven by steady tangential deformation at the surface of the body (so-called squirming motion). In the case of a spherical squirmer, we show that the swimming velocity is systematically less than that in a Newtonian fluid, with a minimum occurring for Weissenberg numbers of…
The distribution of forces on the surface of complex, deforming geometries is an invaluable output of flow simulations. One particular example of such geometries involves self-propelled swimmers. Surface forces can provide significant information about the flow field sensed by the swimmers, and are difficult to obtain experimentally. At the same time, simulations of flow around complex, deforming shapes can be computationally prohibitive when body-fitted grids are used. Alternatively, such simulations may employ penalization techniques. Penalization methods rely on simple Cartesian grids to di…
Trajectories and conformations of uni- and multiflagellar bacteria are studied with a coarse-grained model of a cell comprised of elastic flagella connected to a cell body. The elasticities of both the hook protein (connecting cell body and flagellum) and flagella are varied. Flexibility plays contrasting roles for uni- and multiflagellar swimmers. For a uniflagellar swimmer, hook and/or flagellar buckling occurs above a critical flexibility relative to the torque exerted by the flagellar motor. Addition of a second flagellum greatly expands the parameter regime of stable locomotion, because f…
Swimming microorganisms often self propel in fluids with complex rheology. While past theoretical work indicates that fluid viscoelasticity should hinder their locomotion, recent experiments on waving swimmers suggest a possible non-Newtonian enhancement of locomotion. We suggest a physical mechanism, based on fluid-structure interaction, leading to swimming in a viscoelastic fluid at a higher speed than in a Newtonian one. Using Taylor's two-dimensional swimming sheet model, we solve for the shape of an active swimmer as a balance between the external fluid stresses, the internal driving mome…
Cell motility in viscous fluids is ubiquitous and affects many biological processes, including reproduction, infection, and the marine life ecosystem. Here we review the biophysical and mechanical principles of locomotion at the small scales relevant to cell swimming (tens of microns and below). The focus is on the fundamental flow physics phenomena occurring in this inertia-less realm, and the emphasis is on the simple physical picture. We review the basic properties of flows at low Reynolds number, paying special attention to aspects most relevant for swimming, such as resistance matrices fo…
Undulatory swimming is a widespread propulsion strategy adopted by many small-scale organisms including various single-cell eukaryotes and nematodes. In this work, we report a comprehensive study of undulatory locomotion of a finite filament using (i) approximate resistive force theory (RFT) assuming a local nature of hydrodynamic interaction between the filament and the surrounding viscous liquid, and (ii) particle-based numerical computations taking into account the intra-filament hydrodynamic interaction. Using the ubiquitous model of a propagating sinusoidal waveform, we identify the limit…
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…
An axisymmetric squirmer in a Bingham viscoplastic fluid is studied numerically to determine the effect of a yield stress environment on locomotion. The nonlinearity of the governing equations necessitates numerical methods, which is accomplished by solving a variable-viscosity Stokes equation with a Finite Element approach. The effects of stroke modes, both pure and combined, are investigated and it is found that for the treadmill or "neutral" mode, the swimmer in a yield stress fluid has a lower swimming velocity and uses more power. However, the efficiency of swimming reaches its maximum at…
The locomotion and design of microswimmers are topical issues of current fundamental and applied research. In addition to numerous living and artificial active microswimmers, a passive microswimmer was identified only recently: a soft, Lambda-shaped, non-buoyant particle propagates in a shaken liquid of zero-mean velocity [Jo et al. Phys. Rev. E 94, 063116 (2016)]. We show that this novel passive locomotion mechanism works for realistic non-buoyant, asymmetric Janus microcapsules as well. According to our analytical approximation, this locomotion requires a symmetry breaking caused by differen…
Micro-organisms expend energy moving through complex media. While propulsion speed is an important property of locomotion, efficiency is another factor that may determine the swimming gait adopted by a micro-organism in order to locomote in an energetically favorable manner. The efficiency of swimming in a Newtonian fluid is well characterized for different biological and artificial swimmers. However, these swimmers often encounter biological fluids displaying shear-thinning viscosities. Little is known about how this nonlinear rheology influences the efficiency of locomotion. Does the shear-t…
Micro-scale swimming robots have been envisaged for many medical applications such as targeted drug delivery, where the microrobot will be expected to navigate in a fluid through channels carrying a payload. Alternatively, in many cases, such a payload does not have to be physically bound to the swimmer, but may be instead manipulated and steered through the channel by the microrobot. We investigate this problem of contactless manipulation of a microparticle by mobile microswimmer in a fluid at low Reynolds number. We consider a model of a magnetically actuated artificial microswimmer, whose l…
Synthetic microswimmers show great promise in biomedical applications such as drug delivery and microsurgery. Their locomotion, however, is subject to stringent constraints due to the dominance of viscous over inertial forces at low Reynolds number (Re) in the microscopic world. Furthermore, locomotory gaits designed for one medium may become ineffective in a different medium. Successful biomedical applications of synthetic microswimmers rely on their ability to traverse biological environments with vastly different properties. Here we leverage the prowess of machine learning to present an alt…
Undulatory locomotion is ubiquitous in nature and observed in different media, from the swimming of flagellated microorganisms in biological fluids, to the slithering of snakes on land, or the locomotion of sandfish lizards in sand. Despite the similarity in the undulating pattern, the swimming characteristics depend on the rheological properties of different media. Analysis of locomotion in granular materials is relatively less developed compared with fluids partially due to a lack of validated force models but recently a resistive force theory in granular media has been proposed and shown us…
Combining geometric mechanics theory, laboratory robotic experiment and numerical simulation, we study the locomotion in granular media (GM) of the simplest non-inertial swimmer, the Purcell three-link swimmer. Using granular resistive force laws as inputs, the theory relates translation and rotation of the body to shape changes (movements of the links). This allows analysis, visualization, and prediction of effective movements that are verified by experiment. The geometric approach also facilitates comparison between swimming in GM and in viscous fluids.
Universidad del Valle（山谷大学）；Universidade de São Paulo（圣保罗大学）；Ostschweizer Fachhochschule（东瑞士应用科学大学）
The coupled problem of hydrodynamics and solute transport for the Najafi-Golestanian three-sphere swimmer is studied, with the Reynolds number set to zero and Péclet numbers (Pe) ranging from 0.06 to 60. The adopted method is the numerical simulation of the problem with a finite element code based upon the FEniCS library. For the swimmer executing the optimal locomotion gait, we report the Sherwood number as a function of Pe in homogeneous fluids and confirm that little gain in solute flux is achieved by swimming unless Pe is significantly larger than 10. We also consider the swimmer as an lea…
Swimming at low Reynolds number in Newtonian fluids is only possible through non-reciprocal body deformations due to the kinematic reversibility of the Stokes equations. We consider here a model swimmer consisting of two linked spheres, wherein one sphere is rigid and the other an incompressible neo-Hookean solid. The two spheres are connected by a rod which changes its length periodically. We show that the deformations of the body are non-reciprocal despite the reversible actuation and hence, the elastic two-sphere swimmer propels forward. Our results indicate that even weak elastic deformati…
Most classical work on the hydrodynamics of low-Reynolds-number swimming addresses deterministic locomotion in quiescent environments. Thermal fluctuations in fluids are known to lead to a Brownian loss of the swimming direction. As most cells or synthetic swimmers are immersed in external flows, we consider theoretically in this paper the stochastic dynamics of a model active particle (a self-propelled sphere) in a steady general linear flow. The stochasticity arises both from translational diffusion in physical space, and from a combination of rotary diffusion and run-and-tumble dynamics in …
In this communication we address some hydrodynamic aspects of recently revisited drift mechanism of biogenic mixing in the ocean (Katija and Dabiri, Nature vol. 460, pp. 624-626, 2009). The relevance of the locomotion gait at various spatial scales with respect to the drift is discussed. A hydrodynamic scenario of the drift based on unsteady inertial propulsion, typical for most small marine organisms, is proposed. We estimate its effectiveness by taking into account interaction of a swimmer with the turbulent marine environment. Simple scaling arguments are derived to estimate the comparative…
A growing body of work aims at designing and testing micron-scale synthetic swimmers. One method, inspired by the locomotion of flagellated bacteria, consists of applying a rotating magnetic field to a rigid, helically-shaped, propeller attached to a magnetic head. When the resulting device, termed an artificial bacteria flagellum, is aligned perpendicularly to the applied field, the helix rotates and the swimmer moves forward. Experimental investigation of artificial bacteria flagella shows that at low frequency of the applied field, the axis of the helix does not align perpendicularly to the…
Simulations of undulatory swimming in viscoelastic fluids with large amplitude gaits show concentration of polymer elastic stress at the tips of the swimmers.We use a series of related theoretical investigations to probe the origin of these concentrated stresses. First the polymer stress is computed analytically at a given oscillating extensional stagnation point in a viscoelastic fluid. The analysis identifies a Deborah number (De) dependent Weissenberg number (Wi) transition below which the stress is linear in Wi, and above which the stress grows exponentially in Wi. Next, stress and velocit…
ETH Zurich（苏黎世联邦理工学院）；ETH AI Center（苏黎世联邦理工学院人工智能中心）；Institute of Neuroinformatics, ETH Zurich（苏黎世联邦理工学院神经信息学研究所）
Aquatic locomotion is a classic fluid-structure interaction (FSI) problem of interest to biologists and engineers. Solving the fully coupled FSI equations for incompressible Navier-Stokes and finite elasticity is computationally expensive. Optimizing robotic swimmer design within such a system generally involves cumbersome, gradient-free procedures on top of the already costly simulation. To address this challenge we present a novel, fully differentiable hybrid approach to FSI that combines a 2D direct numerical simulation for the deformable solid structure of the swimmer and a physics-constra…
To explore and react to their environment, living micro-swimmers have developed sophisticated strategies for locomotion - in particular, motility with multiple gaits. To understand the physical principles associated with such a behavioural variability,synthetic model systems capable of mimicking it are needed. Here, we demonstrate bimodal gait switching in autophoretic droplet swimmers. This minimal experimental system is isotropic at rest, a symmetry that can be spontaneously broken due to the nonlinear coupling between hydrodynamic and chemical fields, inducing a variety of flow patterns tha…
The ciliary locomotion and feeding of an axisymmetric micro-swimmer in a complex fluid whose viscosity depends on nutrient concentration are investigated numerically. The micro-swimmer is modeled as having spheroidal geometry, and ciliary beating is modeled by a slip velocity; i.e. a squirmer is adapted. Looking at the coupling between swimming and feeding of spheroidal squirmers, it is found that swimming speed and feeding are most affected by a non-uniform viscosity environment when the ratio of advection-forces to diffusion-transport, characterized by the nondimensional Peclét number, is mo…
We use numerical simulations to address locomotion at zero Reynolds number in viscoelastic (Giesekus) fluids. The swimmers are assumed to be spherical, to self-propel using tangential surface deformation, and the computations are implemented using a finite element method. The emphasis of the study is on the change of the swimming kinematics, energetics, and flow disturbance from Newtonian to viscoelastic, and on the distinction between pusher and puller swimmers. In all cases, the viscoelastic swimming speed is below the Newtonian one, with a minimum obtained for intermediate values of the Wei…
Wake sensing for bioinspired robotic swimmers has been the focus of much investigation owing to its relevance to locomotion control, especially in the context of schooling and target following. Many successful wake sensing strategies have been devised based on models of von Karman-type wakes; however, such wake sensing technologies are invalid in the context of exotic wake types that commonly arise in swimming locomotion. Indeed, exotic wakes can exhibit markedly different dynamics, and so must be modeled and sensed accordingly. Here, we propose a general wake detection protocol for distinguis…
Geometric confinements are frequently encountered in the biological world and strongly affect the stability, topology, and transport properties of active suspensions in viscous flow. Based on a far-field analytical model, the low-Reynolds-number locomotion of a self-propelled microswimmer moving inside a clean viscous drop or a drop covered with a homogeneously distributed surfactant, is theoretically examined. The interfacial viscous stresses induced by the surfactant are described by the well-established Boussinesq-Scriven constitutive rheological model. Moreover, the active agent is represe…
