In this paper we study swimming of a model organism, the so-called Taylor's swimming sheet, in a viscoelastic fluid close to a solid boundary. This situation comprises natural habitats of many swimming microorganisms, and while previous investigations have considered the effects of both swimming next to a boundary and swimming in a viscoelastic fluid, seldom have both effects been considered simultaneously. We re-visit the small wave amplitude result obtained by Elfring and Lauga (Gwynn J. Elfring and Eric Lauga, in Saverio E. Spagnolie, editor, Complex Fluids in Biological Systems, Springer New York, New York, NY, 2015), and give a mechanistic explanation to the decoupling of the effects of viscoelasticity, which tend to slow the sheet, and the presence of the boundary, which tends to speed up the sheet. We also develop a numerical spectral method capable of finding the swimming speed of a waving sheet with an arbitrary amplitude and waveform. We use it to show that the decoupling mentioned above does not hold at finite wave amplitudes and that for some parameters the presence of a boundary can cause the viscoelastic effects to increase the swimming speed of microorganisms.
We present a realization of a fast interfacial Marangoni microswimmer by a half-spherical alginate capsule at the air-water interface, which diffusively releases water-soluble spreading molecules (weak surfactants such as polyethylene glycol (PEG)), which act as "fuel" by modulating the air-water interfacial tension. For a number of different fuels, we can observe symmetry breaking and spontaneous propulsion although the alginate particle and emission are isotropic. The propulsion mechanism is similar to soap or camphor boats, which are, however, typically asymmetric in shape or emission to se…
Studies of model microswimmers have significantly contributed to the understanding of the principles of self-propulsion we have today. However, only a small number of microswimmer types have been amenable to analytic modeling, and further development of such approaches is necessary to identify the key features of these active systems. Here we present a general perturbative calculation scheme for swimmers composed of beads interacting by harmonic potentials, driven by an arbitrary force protocol. The hydrodynamic interactions are treated using the Oseen and Rotne-Pragner approximations. We vali…
The current work studies the dynamics of a microswimmer in pressure-driven flow of a weakly viscoelastic fluid. Employing the second-order fluid model, we show that the self-propelling swimmer experiences a viscoelastic swimming lift in addition to the well-known passive lift that arises from its resistance to shear flow. Using the reciprocal theorem, we evaluate analytical expressions for the swimming lift experienced by neutral and pusher/puller-type swimmers and show that they depend on the hydrodynamic signature associated with the swimming mechanism. We find that for neutral swimmers focu…
We propose two-dimensional organic poly(heptazine imide) (PHI) carbon nitride microparticles as light-driven microswimmers in various ionic and biological media. Their demonstrated high-speed (15-23 $μ$m/s) swimming in multi-component ionic solutions with concentrations up to 1 M and without dedicated fuels is unprecedented, overcoming one of the bottlenecks of previous light-driven microswimmers. Such high ion tolerance is attributed to a favorable interplay between the particle's textural and structural nanoporosity and optoionic properties, facilitating ionic interactions in solutions with …
In this paper we are interested in optimizing the shape of multi-flagellated helical microswimmers. Mimicking the propagation of helical waves along the flagella, they self-propel by rotating their tails. The swimmer's dynamics is computed using the Boundary Element Method, implemented in the open source Matlab library $Gypsilab$. We exploit a Bayesian optimization algorithm to maximize the swimmer's speeds through their shape optimization. Our results show that the optimal tail shapes are helices with large wavelength, such that the shape periodicity is disregarded. Moreover, the best propuls…
Viscoelastic fluids impact the locomotion of swimming microorganisms and can be harnessed to devise new types of self-propelling devices. Here we report on experiments demonstrating the use of normal stress differences for propulsion. Rigid dumbbells are rotated by an external magnetic field along their axis of symmetry in a Boger fluid. When the dumbbell is asymmetric (snowman geometry), non-Newtonian normal stress differences lead to net propulsion in the direction of the smaller sphere. The use of a simple model allows to rationalise the experimental results and to predict the dependence of…
Self-propelling particles or microswimmers have opened a new field of investigation with both fundamental and practical perspectives. They represent very convenient model objects for experimental studies of active matter, and have implications in nano-robotics, drug delivery, and more. Here, we summarize recent advances in theoretical description of the self-propulsion of catalytic microswimmers that non-uniformly release ions, including its physical origins and the current switch in focus to non-linear effects, geometric tuning and more. In particular, we show that the ionic self-propulsion a…
The motion of microswimmers in complex flows is ruled by the interplay between swimmer propulsion and the dynamics induced by the fluid velocity field. Here we study the motion of a chiral microswimmer whose propulsion is provided by the spinning of a helical tail with respect to its body in a simple shear flow. Thanks to an efficient computational strategy that allowed us to simulate thousands of different trajectories, we show that the tail shape dramatically affects the swimmer's motion. In the shear dominated regime, the swimmers carrying an elliptical helical tail show several different J…
We study swimming of small spherical particles who regulate fluid flow on their surface by applying tangential squirming strokes. We derive translational and rotational velocities for any given stroke which is not restricted by axial symmetry as assumed usually. The formulation includes inertia of both the fluid and the swimmer, motivated by inertia's relevance for large Volvox colonies. We show that inertial contribution to mean speed comes from dynamic coupling between translation and rotation, which occurs only for strokes that break axial symmetry. Remarkably, this effect enables overcomin…
Microswimmers create bicontinuous emulsions in binary fluids
Indian Institute of Technology Kharagpur（印度理工学院卡拉格普尔分校）；EMBL Barcelona（欧洲分子生物学实验室巴塞罗那分部）；Barcelona Collaboratorium for Modelling and Predictive Biology（巴塞罗那建模与预测生物合作组织）；Univ. Bordeaux（波尔多大学）
Synchronization of two actuated sheets serves as a simple model for the interaction between flagellated microswimmers. Various factors, including inertia, sheet elasticity, and fluid viscoelasticity, have been suggested to facilitate the synchronization of two sheets; however, the importance of different contributions to this process still remains unclear. We perform a systematic investigation of competing effects of inertia, sheet elasticity, and fluid viscoelasticity on the synchronization of two sheets. Characteristic time $τ^\mathrm{s}$ for the synchronization caused by inertial effects is…
In this work, we numerically investigate the dynamics of a self-propelling autophoretic Janus particle in a weakly viscoelastic fluid. The self-propulsion is achieved by an asymmetry in the properties of the surface of the Janus particle that drives a surface slip velocity and bulk flow. Here we investigate the effect of viscoelasticity on this advection-diffusion problem over a range of Péclet and Damköhler numbers. Particles are found to swim faster, or slower, in viscoelastic fluids, and we show how reaction and diffusion rates affect the viscoelastic stresses that lead to changes in propul…
In the study of microscopic flows, self-propulsion has been particularly topical in recent years, with the rise of miniature artificial swimmers as a new tool for flow control, low Reynolds number mixing, micromanipulation or even drug delivery. It is possible to take advantage of interfacial physics to propel these micro-robots, as demonstrated by recent experiments using the proximity of an interface, or the interface itself, to generate propulsion at low Reynolds number. This paper discusses how a nearby interface can provide the symmetry breaking necessary for propulsion. An overview of re…
The effects of fluid elasticity on the swimming behavior of the nematode \emph{Caenorhabditis elegans} are experimentally investigated by tracking the nematode's motion and measuring the corresponding velocity fields. We find that fluid elasticity hinders self-propulsion. Compared to Newtonian solutions, fluid elasticity leads to 35% slower propulsion speed. Furthermore, self-propulsion decreases as elastic stresses grow in magnitude in the fluid. This decrease in self-propulsion in viscoelastic fluids is related to the stretching of flexible molecules near hyperbolic points in the flow.
Synthetic microswimmers offer models for cell motility and their tunability makes them promising candidates for biomedical applications. Here we measure the acoustic propulsion of bimetallic micro-rods that, when trapped at the nodal plane of a MHz acoustic resonator, swim with speeds of up to 300 microns per second. While past acoustic streaming models predict speeds that are more than one order of magnitude smaller than our measurements, we demonstrate that the acoustic locomotion of the rods is driven by a viscous, non-reciprocal mechanism relying on shape anisotropy akin to that used by sw…
Flagella beating in complex fluids are significantly influenced by viscoelastic stresses. Relevant examples include the ciliary transport of respiratory airway mucus and the motion of spermatozoa in the mucus-filled female reproductive tract. We consider the simplest model of such propulsion and transport in a complex fluid, a waving sheet of small amplitude free to move in a polymeric fluid with a single relaxation time. We show that, compared to self-propulsion in a Newtonian fluid occurring at a velocity U_N, the sheet swims (or transports fluid) with velocity U / U_N = [1+De^2 (eta_s)/(eta…
The ability of microswimmers to deploy optimal propulsion strategies is of paramount importance for their locomotory performance and survival at low Reynolds numbers. Although for perfectly spherical swimmers minimum dissipation requires a neutral type swimming, any departure from the spherical shape may lead the swimmer to adopt a new propulsion strategy, namely those of puller- or pusher-type swimming. In this study, by using the minimum dissipation theorem for microswimmers, we determine the flow field of an optimal nearly spherical swimmer, and show that indeed depending on the shape profi…
Most biological fluids are viscoelastic, meaning that they have elastic properties in addition to the dissipative properties found in Newtonian fluids. Computational models can help us understand viscoelastic flow, but are often limited in how they deal with complex flow geometries and suspended particles. Here, we present a lattice Boltzmann solver for Oldroyd-B fluids that can handle arbitrarily-shaped fixed and moving boundary conditions, which makes it ideally suited for the simulation of confined colloidal suspensions. We validate our method using several standard rheological setups, and …
For natural microswimmers, the interplay of swimming activity and external flow can promote robust motion, e.g. propulsion against ("upstream rheotaxis") or perpendicular to the direction of flow. These effects are generally attributed to their complex body shapes and flagellar beat patterns. Here, using catalytic Janus particles as a model experimental system, we report on a strong directional response that occurs for spherical active particles in a channel flow. The particles align their propulsion axes to be nearly perpendicular to both the direction of flow and the normal vector of a nearb…
Self-propelled particles can exhibit surprising non-equilibrium behaviors, and how they interact with obstacles or boundaries remains an important open problem. Here we show that chemically propelled micro-rods can be captured, with little change in their speed, into close orbits around solid spheres resting on or near a horizontal plane. We show that this interaction between sphere and particle is short-range, occurring even for spheres smaller than the particle length, and for a variety of sphere materials. We consider a simple model, based on lubrication theory, of a force- and torque-free …
Many microorganisms swim in fluids with complex rheological properties. Although much is now understood about motion of these swimmers in Newtonian fluids, the understanding is still developing in non-Newtonian fluids --- this understanding is crucial for various biomimetic and biomedical applications. Here we study a common model for microswimmers, the squirmer model, in two common viscoelastic fluid models, the Giesekus fluid model and fluids of differential type (grade three), at zero Reynolds number. Through this article we address a recent commentary that discussed suitable values of para…
Self-propulsion at low Reynolds number is notoriously restricted, a concept that is commonly known as the "scallop theorem". Here we present a truly self-propelled swimmer (force- and torque- free) that, while unable to swim in a Newtonian fluid due to the scallop theorem, propels itself in a non-Newtonian fluid as a result of fluid elasticity. This propulsion mechanism is demonstrated using a robotic swimmer, comprised of a "head" sphere and a "tail" sphere, whose swimming speed is shown to have reasonable agreement with a microhydrodynamic asymptotic theory and numerical simulations. Schlier…
Several recent experiments investigate the orientational and transport behavior of self-driven bacteria and colloidal particles in nematic liquid crystals. Correspondingly, we study theoretically the dynamics of a minimal model microswimmer in a uniaxially anisotropic fluid. As a first step, the hydrodynamic Green's function providing the resulting fluid flow in response to a localized force acting on the anisotropic fluid is derived analytically. On this basis, the behavior of both puller- and pusher-type microswimmers in the anisotropic fluid is analyzed. Depending on the propulsion mechanis…
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…
Many practically relevant materials combine properties of viscous fluids and elastic solids to viscoelastic behavior. Our focus is on the induced dynamic behavior of damped finite-sized particulate inclusions in such substances. We explicitly describe history-dependent interactions that emerge between the embedded particles. These interactions are mediated by the viscoelastic surroundings. They result from the flows and distortions of the viscoelastic medium when induced by the rigid inclusions. Both, viscoelastic environments of terminal fluid-like flow or of completely reversible damped elas…
The problem of optimal microscopic swimming in a noisy environment is analyzed. A simplified model in which propulsion is generated by the relative motion of three spheres connected by immaterial links has been considered. We show that an optimized noisy microswimmer requires less power for propulsion (on the average), than an optimal noiseless counterpart, migrating with identical mean velocity and swimming stroke amplitude. We also show that noise can be used to overcome some of the limitations of the scallop theorem, and have a swimmer that is able to propel itself with control over just on…
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…
Propulsion through fluids is a key component in the life cycle of many microbes, be it in development, infection, or simply finding nutrients. In systems of biomedical relevance, this propulsion is often through polymer suspensions that endow the fluid with complex non-Newtonian properties, such as shear-thinning and viscoelastic behaviour. Due to the complexity of 3D non-Newtonian modelling, 2D undulatory propulsion has recently been extensively-studied as a means of garnering physical intuition for these systems. However, while streamlines, swimming speeds, and swimmer trajectories are strik…
Active matter systems such as eukaryotic cells and bacteria continuously transform chemical energy to motion. Hence living systems exert active stresses on the complex environments in which they reside. One recurring aspect of this complexity is the viscoelasticity of the medium surrounding living systems: bacteria secrete their own viscoelastic extracellular matrix, and cells constantly deform, proliferate, and self-propel within viscoelastic networks of collagen. It is therefore imperative to understand how active matter modifies, and gets modified by, viscoelastic fluids. Here, we present a…
Interaction of motile microrganisms with a nearby solid substrate is a well studied phenomenon. However, the effects of hydrodynamic slippage on the substrate have received a little attention. In the present study, within the framework of the squirmer model, we impose a tangential velocity at the swimmer surface as a representation of the ciliatory propulsion and subsequently obtain exact solution of the Stokes equation based on a combined analytical-numerical approach. We illustrate how the near-wall swimming velocities are non-trivially altered by the interaction of wall slip and hydrodynami…
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 small organisms self-propel in viscous fluids using travelling wave-like deformation of their bodies or appendages. Examples include small nematodes moving through soil using whole-body undulations or spermatozoa swimming through mucus using flagellar waves. When self-propulsion occurs in a non-Newtonian fluid, one fundamental question is whether locomotion will occur faster or slower than in a Newtonian environment. Here we consider the general problem of swimming using small-amplitude periodic waves in a viscoelastic fluid described by the classical Oldroyd-B constitutive relationship. …
In a variety of biological situations, swimming cells have to move through complex fluids. Similarly, mucociliary clearance involves the transport of polymeric fluids by beating cilia. Here, we consider the extent to which complex fluids could be exploited for force generation on small scales. We consider a prototypical reciprocal motion (i.e. identical under time-reversal symmetry): The periodic flapping of a tethered semi-infinite plane. In the Newtonian limit, such motion cannot be used for force generation according to Purcell's scallop theorem. In a polymeric fluid (Oldroyd-B, and its gen…
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…
Shear-thinning is an important rheological property of many biological fluids, such as mucus, whereby the apparent viscosity of the fluid decreases with shear. Certain microscopic swimmers have been shown to progress more rapidly through shear-thinning fluids, but is this behavior generic to all microscopic swimmers, and what are the physics through which shear-thinning rheology affects a swimmer's propulsion? We examine swimmers employing prescribed stroke kinematics in two-dimensional, inertialess Carreau fluid: shear-thinning "Generalized Stokes" flow. Swimmers are modeled, using the method…
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 …
