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Shear-thinning viscosity is a non-Newtonian behaviour that active particles often encounter in biological fluids such as blood and mucus. The fundamental question of how this ubiquitous non-Newtonian rheology affects the propulsion of active particles has attracted substantial interest. In particular, spherical Janus particles driven by self-diffusiophresis, a major physico-chemical propulsion mechanism of synthetic active particles, were shown to always swim slower in a shear-thinning fluid than in a Newtonian fluid. In this work, we move beyond the spherical limit to examine the effect of particle eccentricity on self-diffusiophoretic propulsion in a shear-thinning fluid. We use a combination of asymptotic analysis and numerical simulations to show that shear-thinning rheology can enhance self-diffusiophoretic propulsion of a spheroidal particle, in stark contrast to previous findings for the spherical case. A systematic characterization of the dependence of the propulsion speed on the particle's active surface coverage has also uncovered an intriguing feature associated with the propulsion speeds of a pair of complementarily coated particles not previously reported. Symmetry arguments are presented to elucidate how this new feature emerges as a combined effect of anisotropy of the spheroidal geometry and nonlinearity in fluid rheology., Comment: 21 Pages, 8 figures

Microorganisms and synthetic microswimmers often encounter complex environments consisting of networks of obstacles embedded into viscous fluids. Such settings include biological media, such as mucus with filamentous networks, as well as environmental scenarios, including wet soil and aquifers. A fundamental question in studying their locomotion is how the impermeability of these porous media impact their propulsion performance compared with the case that in a purely viscous fluid. Previous studies showed that the additional resistance due to the embedded obstacles leads to an enhanced propulsion of different types of swimmers, including undulatory swimmers, helical swimmers, and squirmers. In this work we employ a canonical three-sphere swimmer model to probe the impact of propulsion in porous media. The Brinkman equation is utilized to model a sparse network of stationary obstacles embedded into an incompressible Newtonian liquid. We present both a far-field theory and numerical simulations to characterize the propulsion performance of the swimmer in such porous media. In contrast to enhanced propulsion observed in other swimmer models, our results reveal that both the propulsion speed and efficiency of the three-sphere swimmer are largely reduced by the impermeability of the porous medium. We attribute the substantial reduction in propulsion performance to the screened hydrodynamic interactions among the spheres due to the more rapid spatial decays of flows in Brinkman media. These results highlight how enhanced or hindered propulsion in porous media is largely dependent on individual propulsion mechanisms. The specific example and physical insights provided here may guide the design of synthetic microswimmers for effective locomotion in porous media in their potential biological and environmental applications., Comment: 23 pages, 4 figures

Swimming at the microscale has recently garnered substantial attention due to the fundamental biological significance of swimming microorganisms and the wide range of biomedical applications for artificial microswimmers. These microswimmers invariably find themselves surrounded by different confining boundaries, which can impact their locomotion in significant and diverse ways. In this work, we employ a widely used three-sphere swimmer model to investigate the effect of confinement on swimming at low Reynolds numbers. We conduct theoretical analysis via the point-particle approximation and numerical simulations based on the finite element method to examine the motion of the swimmer along the centerline in a capillary tube. The axisymmetric configuration reduces the motion to one-dimensional movement, which allows us to quantify how the degree of confinement affects the propulsion speed in a simple manner. Our results show that the confinement does not significantly affect the propulsion speed until the ratio of the radius of the tube to the radius of the sphere is in the range of $\mathcal{O}(1)-\mathcal{O}(10)$, where the swimmer undergoes substantial reduction in its propulsion speed as the radius of the tube decreases. We provide some physical insights into how reduced hydrodynamic interactions between moving spheres under confinement may hinder the propulsion of the three-sphere swimmer. We also remark that the reduced propulsion performance stands in stark contrast to the enhanced helical propulsion observed in a capillary tube, highlighting how the manifestation of confinement effects can vary qualitatively depending on the propulsion mechanisms employed by the swimmers., Comment: 10 pages, 7 figures, revised manuscript resubmitted to Phys Fluids

The effects of viscoelasticity have been shown to manifest themselves via symmetry breaking. In this investigation, we show a novel phenomenon that arises from this idea. We observe that when a dense sphere is rotated near a wall (the rotation being aligned with the wall-normal direction and gravity), it levitates to a fixed distance away from the wall. Since the shear is larger in the gap (between the sphere and the wall) than in the open side of the sphere, the shear-induced elastic stresses are thus asymmetric, resulting in a net elastic vertical force that balances the weight of the sphere. We conduct experiments, theoretic models, and numerical simulations for rotating spheres of various sizes and densities in a Boger-type fluid. In the small Deborah number range, the results are collapsed into a universal trend by considering a dimensionless group of the ratio of elastic to gravitational forces.

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 alternative approach to designing low Re swimmers. Instead of specifying any locomotory gaits \textit{a priori}, here a swimmer develops its own propulsion strategy based on its interactions with the surrounding medium via reinforcement learning. This self-learning capability enables the swimmer to modify its propulsion strategy in response to different environments. We illustrate this new approach using a minimal example that integrates a standard reinforcement learning algorithm ($Q$-learning) into the locomotion of a swimmer consisting of an assembly of spheres connected by extensible rods. We showcase theoretically that this first self-learning swimmer can recover a previously known propulsion strategy without prior knowledge in low Re locomotion, identify more effective locomotory gaits when the number of spheres increases, and adapt its locomotory gaits in different media. These results represent initial steps towards the design of a new class of self-learning, adaptive (or "smart") swimmers with robust locomotive capabilities to traverse complex biological environments.

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-thinning rheology render swimming more efficient or less? How does the swimming efficiency depend on the propulsion mechanism of a swimmer and rheological properties of the surrounding shear-thinning fluid? In this work, we address these fundamental questions on the efficiency of locomotion in a shear-thinning fluid by considering the squirmer model as a general locomotion model to represent different types of swimmers. Our analysis reveals how the choice of surface velocity distribution on a squirmer may reduce or enhance the swimming efficiency. We determine optimal shear rates at which the swimming efficiency can be substantially enhanced com- pared with the Newtonian case. The non-trivial variations of swimming efficiency prompt questions on how micro-organisms may tune their swimming gaits to exploit the shear-thinning rheology. The findings also provide insights into how artificial swimmers should be designed to move through complex media efficiently.

Biological locomotion in nature is often achieved by the interaction between a flexible body and its surrounding medium. The interaction of a flexible body with granular media is less understood compared with viscous fluids partially due to its complex rheological properties. In this work we explore the effect of flexibility on granular propulsion by considering a simple mechanical model where a rigid rod is connected to a torsional spring that is under a displacement actuation using a granular resistive force theory. Through a combined numerical and asymptotic investigation, we characterize the propulsive dynamics of such a flexible flapper in relation to the actuation amplitude and spring stiffness and compare these dynamics with those observed in a viscous fluid. In addition, we demonstrate that the maximum possible propulsive force can be obtained in the steady propulsion limit with a finite spring stiffness and large actuation amplitude. These results may apply to the development of synthetic locomotive systems that exploit flexibility to move through complex terrestrial media., Comment: 9 pages, 5 figures

At low Reynolds numbers the locomotive capability of a body can be dramatically hindered by the absence of inertia. In this work, we show how propulsive performance in this regime can be significantly enhanced by employing spatially varying flexibility. As a prototypical example, we consider the propulsive thrust generated by a filament periodically driven at one end. The rigid case leads to zero propulsion, as so constrained by Purcell's scallop theorem, while for uniform filaments there exists a bending stiffness maximizing the propulsive force at a given frequency; here we demonstrate explicitly how considerable further improvement (greater than 37% enhancement compared with the optimal uniform stiffness) can be achieved by simply varying the stiffness along the filament. The optimal flexibility distribution is strongly configuration-dependent: while increasing the flexibility towards the tail-end enhances the propulsion of a clamped filament, for a hinged filament decreasing the flexibility towards the tail-end is instead favorable. The results reveal new design principles for maximizing propulsion at low Reynolds numbers, potentially useful for developing synthetic micro-swimmers requiring large propulsive force for various biomedical applications., Comment: 5 figures

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 useful in studying the locomotion of a sand-swimming lizard. Here we employ the proposed model to investigate the swimming characteristics of a slender filament, of both finite and infinite length, undulating in a granular medium and compare the results with swimming in viscous fluids. In particular, we characterize the effects of drifting and pitching in terms of propulsion speed and efficiency for a finite sinusoidal swimmer. We also find that, similar to Lighthill's results using resistive force theory in viscous fluids, the sawtooth swimmer is the optimal waveform for propulsion speed at a given power consumption in granular media. The results complement our understanding of undulatory locomotion and provide insights into the effective design of locomotive systems in granular media.

Many microorganisms find themselves immersed in fluids displaying non-Newtonian rheological properties such as viscoelasticity and shear-thinning viscosity. The effects of viscoelasticity on swimming at low Reynolds numbers have already received considerable attention, but much less is known about swimming in shear-thinning fluids. A general understanding of the fundamental question of how shear-thinning rheology influences swimming still remains elusive. To probe this question further, we study a spherical squirmer in a shear-thinning fluid using a combination of asymptotic analysis and numerical simulations. Shear-thinning rheology is found to affect a squirming swimmer in nontrivial and surprising ways; we predict and show instances of both faster and slower swimming depending on the surface actuation of the squirmer. We also illustrate that while a drag and thrust decomposition can provide insights into swimming in Newtonian fluids, extending this intuition to problems in complex media can prove problematic.

Experiments on the acoustic vibrations of elastic nanostructures in fluid media have been used to study the mechanical properties of materials, as well as for mechanical and biological sensing. The medium surrounding the nanostructure is typically modeled as a Newtonian fluid. A recent experiment however suggested that high-frequency longitudinal vibration of bipyramidal nanoparticles could trigger a viscoelastic response in water-glycerol mixtures [M. Pelton et al., "Viscoelastic flows in simple liquids generated by vibrating nanostructures," Phys. Rev. Lett. 111, 244502 (2013)]. Motivated by these experimental studies, we first revisit a classical continuum mechanics problem of the purely radial vibration of an elastic sphere, also called the breathing mode, in a compressible viscous fluid, and then extend our analysis to a viscoelastic medium using the Maxwell fluid model. The effects of fluid compressibility and viscoelasticity are discussed. Although in the case of longitudinal vibration of bipyramidal nanoparticles, the effects of fluid compressibility were shown to be negligible, we demonstrate that it plays a significant role in the breathing mode of an elastic sphere. On the other hand, despite the different vibration modes, the breathing mode of a sphere triggers a viscoelastic response in water-glycerol mixtures similar to that triggered by the longitudinal vibration of bipyramidal nanoparticles. We also comment on the effect of fluid viscoelasticity on the idea of destroying virus particles by acoustic resonance.

The locomotion of microorganisms in fluids is ubiquitous and plays an important role in numerous biological processes. In this chapter we present an overview of theoretical modeling for low-Reynolds-number locomotion., Comment: Chapter in "Low-Reynolds-Number Flows: Fluid-Structure Interactions", C. Duprat and H. A. Stone (Eds.), Royal Society of Chemistry Soft Matter Series (2014, to appear)

A number of swimming microorganisms such as ciliates ($\textit{Opalina}$) and multicellular colonies of flagellates ($\textit{Volvox}$) are approximately spherical in shape and swim using beating arrays of cilia or short flagella covering their surfaces. Their physical actuation on the fluid may be mathematically modeled as the generation of surface velocities on a continuous spherical surface -- a model known in the literature as squirming, which has been used to address various aspects of the biological physics of locomotion. Previous analyses of squirming assumed axisymmetric fluid motion and hence restricted all swimming kinematics to take place along a line. In this paper we generalize squirming to three spatial dimensions. We derive analytically the flow field surrounding a spherical squirmer with arbitrary surface motion, and use it to derive its three-dimensional translational and rotational swimming kinematics. We then use our results to physically interpret the flow field induced by the swimmer in terms of fundamental flow singularities up to terms decaying spatially as $\sim 1/r^3$. Our results will enable to develop new models in biological physics, in particular in the area of hydrodynamic interactions and collective locomotion.

Many biological fluids have polymeric microstructures and display non-Newtonian rheology. We take advantage of such nonlinear fluid behavior and combine it with geometrical symmetry-breaking to design a novel small-scale propeller able to move only in complex fluids. Its propulsion characteristics are explored numerically in an Oldroyd-B fluid for finite Deborah numbers while the small Deborah number limit is investigated analytically using a second-order fluid model. We then derive expressions relating the propulsion speed to the rheological properties of the complex fluid, allowing thus to infer the normal stress coefficients in the fluid from the locomotion of the propeller. Our simple mechanism can therefore be used either as a non-Newtonian micro-propeller or as a micro-rheometer.

In addition to conventional planar and helical flagellar waves, insect sperm flagella have also been observed to display a double-wave structure characterized by the presence of two superimposed helical waves. In this paper, we present a hydrodynamic investigation of the locomotion of insect spermatozoa exhibiting the double-wave structure, idealized here as superhelical waves. Resolving the hydrodynamic interactions with a non-local slender body theory, we predict the swimming kinematics of these superhelical swimmers based on experimentally collected geometric and kinematic data. Our consideration provides insight into the relative contributions of the major and minor helical waves to swimming; namely, propulsion is due primarily to the minor wave, with negligible contribution from the major wave. We also explore the dependence of the propulsion speed on geometric and kinematic parameters, revealing counter-intuitive results, particularly for the case when the minor and major helical structures are of opposite chirality.

Micro/nano-scale propulsion has attracted considerable recent attention due to its promise for biomedical applications such as targeted drug delivery. In this paper, we report on a new experimental design and theoretical modelling of high-speed fuel-free magnetically-driven propellers which exploit the flexibility of nanowires for propulsion. These readily prepared nanomotors display both high dimensional propulsion velocities (up to ~ 21 micrometer per second) and dimensionless speeds (in body lengths per revolution) when compared with natural microorganisms and other artificial propellers. Their propulsion characteristics are studied theoretically using an elastohydrodynamic model which takes into account the elasticity of the nanowire and its hydrodynamic interaction with the fluid medium. The critical role of flexibility in this mode of propulsion is illustrated by simple physical arguments, and is quantitatively investigated with the help of an asymptotic analysis for small-amplitude swimming. The theoretical predictions are then compared with experimental measurements and we obtain good agreement. Finally, we demonstrate the operation of these nanomotors in a real biological environment (human serum), emphasizing the robustness of their propulsion performance and their promise for biomedical applications., Comment: Electronic supplementary information (ESI) available: Experimental details and a series of videos demonstrating nanowire swimmers in motion. See DOI: 10.1039/c1sm05503h

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 weak or even absent.

In a world without inertia, Purcell's scallop theorem states that in a Newtonian fluid a time-reversible motion cannot produce any net force or net flow. Here we consider the extent to which the nonlinear rheological behavior of viscoelastic fluids can be exploited to break the constraints of the scallop theorem in the context of fluid pumping. By building on previous work focusing on force generation, we consider a simple, biologically-inspired geometrical example of a flapper in a polymeric (Oldroyd-B) fluid, and calculate asymptotically the time-average net fluid flow produced by the reciprocal flapping motion. The net flow occurs at fourth order in the flapping amplitude, and suggests the possibility of transporting polymeric fluids using reciprocal motion in simple geometries even in the absence of inertia. The induced flow field and pumping performance are characterized and optimized analytically. Our results may be useful in the design of micro-pumps handling complex fluids.

Experimental studies have demonstrated that spermatozoa synchronize their flagella when swimming in close proximity. In a Newtonian fluid, it was shown theoretically that such synchronization arises passively due to hydrodynamic forces between the two swimmers if their waveforms exhibit a front-back geometrical asymmetry. Motivated by the fact that most biological fluids possess a polymeric microstructure, we address here synchronization in a viscoelastic fluid analytically. Using a two-dimensional infinite sheet model we show that the presence of polymeric stresses removes the geometrical asymmetry constraint, and therefore even symmetric swimmers synchronize. Such synchronization occurs on asymptotically faster time scales than in a Newtonian fluid, and the swimmers are seen to be driven into a stable in-phase conformation minimizing the energy dissipated in the surrounding fluid.

Small-scale locomotion plays an important role in biology. Different modelling approaches have been proposed in the past. The simplest model is an infinite inextensible two-dimensional waving sheet, {originally introduced by Taylor}, which serves as an idealized geometrical model for both spermatozoa locomotion and ciliary transport in Stokes flow. Here we complement classic steady-state calculations by deriving the transient low-Reynolds number swimming speed of such a waving sheet when starting from rest (small-amplitude initial-value problem). We also determine the transient fluid flow in the `pumping' setup where the sheet is not free to move but instead generates a net fluid flow around it. The time scales for these two problems, which in general govern transient effects in transport and locomotion at low Reynolds numbers, are also derived using physical arguments.