Propagating density spikes in light-powered motility-ratchets.

Combining experiments and computer simulations, we use a spatially periodic and flashing light-field to direct the motion of phototactic active colloids. Here, the colloids self-organize into a density spike pattern, which resembles a shock wave and propagates over long distances, almost without dispersing. The underlying mechanism involves a synchronization of the colloids with the light-field, so that particles see the same intensity gradient each time the light-pattern is switched on, but no gradient in between (for example). This creates pulsating transport whose strength and direction can be controlled via the flashing protocol and the self-propulsion speed of the colloids. Our results might be useful for drug delivery applications and can be used to segregate active colloids by their speed.

Colloidal Brazil nut effect in microswimmer mixtures induced by motility contrast.

We numerically and experimentally study the segregation dynamics in a binary mixture of microswimmers which move on a two-dimensional substrate in a static periodic triangular-like light intensity field. The motility of the active particles is proportional to the imposed light intensity, and they possess a motility contrast, i.e., the prefactor depends on the species. In addition, the active particles also experience a torque aligning their motion towards the direction of the negative intensity gradient. We find a segregation of active particles near the intensity minima where typically one species is localized close to the minimum and the other one is centered around in an outer shell. For a very strong aligning torque, there is an exact mapping onto an equilibrium system in an effective external potential that is minimal at the intensity minima. This external potential is similar to (height-dependent) gravity such that one can define effective "heaviness" of the self-propelled particles. In analogy to shaken granular matter in gravity, we define a "colloidal Brazil nut effect" if the heavier particles are floating on top of the lighter ones. Using extensive Brownian dynamics simulations, we identify system parameters for the active colloidal Brazil nut effect to occur and explain it based on a generalized Archimedes' principle within the effective equilibrium model: heavy particles are levitated in a dense fluid of lighter particles if their effective mass density is lower than that of the surrounding fluid. We also perform real-space experiments on light-activated self-propelled colloidal mixtures which confirm the theoretical predictions.

Viscotaxis: Microswimmer Navigation in Viscosity Gradients.

The survival of many microorganisms, like Leptospira or Spiroplasma bacteria, can depend on their ability to navigate towards regions of favorable viscosity. While this ability, called viscotaxis, has been observed in several bacterial experiments, the underlying mechanism remains unclear. We provide a framework to study viscotaxis of biological or synthetic self-propelled swimmers in slowly varying viscosity fields and show that suitable body shapes create viscotaxis based on a systematic asymmetry of viscous forces acting on a microswimmer. Our results shed new light on viscotaxis in Spiroplasma and Leptospira and suggest that dynamic body shape changes exhibited by both types of microorganisms may have an unrecognized functionality: to prevent them from drifting to low viscosity regions where they swim poorly. The present theory classifies microswimmers regarding their ability to show viscotaxis and can be used to design synthetic viscotactic swimmers, e.g., for delivering drugs to a target region distinguished by viscosity.

Helical paths, gravitaxis, and separation phenomena for mass-anisotropic self-propelling colloids: Experiment versus theory.

The self-propulsion mechanism of active colloidal particles often generates not only translational but also rotational motion. For particles with an anisotropic mass density under gravity, the motion is usually influenced by a downwards oriented force and an aligning torque. Here we study the trajectories of self-propelled bottom-heavy Janus particles in three spatial dimensions both in experiments and by theory. For a sufficiently large mass anisotropy, the particles typically move along helical trajectories whose axis is oriented either parallel or antiparallel to the direction of gravity (i.e., they show gravitaxis). In contrast, if the mass anisotropy is small and rotational diffusion is dominant, gravitational alignment of the trajectories is not possible. Furthermore, the trajectories depend on the angular self-propulsion velocity of the particles. If this component of the active motion is strong and rotates the direction of translational self-propulsion of the particles, their trajectories have many loops, whereas elongated swimming paths occur if the angular self-propulsion is weak. We show that the observed gravitational alignment mechanism and the dependence of the trajectory shape on the angular self-propulsion can be used to separate active colloidal particles with respect to their mass anisotropy and angular self-propulsion, respectively.

Depolarized light scattering from prolate anisotropic particles: The influence of the particle shape on the field autocorrelation function.

We provide a theoretical analysis for the intermediate scattering function typically measured in depolarized dynamic light scattering experiments. We calculate the field autocorrelation function g1(VH)(Q,t) in dependence on the wave vector Q and the time t explicitly in a vertical-horizontal scattering geometry for differently shaped solids of revolution. The shape of prolate cylinders, spherocylinders, spindles, and double cones with variable aspect ratio is expanded in rotational invariants flm(r). By Fourier transform of these expansion coefficients, a formal multipole expansion of the scattering function is obtained, which is used to calculate the weighting coefficients appearing in the depolarized scattering function. In addition to translational and rotational diffusion, especially the translational-rotational coupling of shape-anisotropic objects is considered. From the short-time behavior of the intermediate scattering function, the first cumulants Γ(Q) are calculated. In a depolarized scattering experiment, they deviate from the simple proportionality to Q(2). The coefficients flm(Q) strongly depend on the geometry and aspect ratio of the particles. The time dependence, in addition, is governed by the translational and rotational diffusion tensors, which are calculated by means of bead models for differently shaped particles in dependence on their aspect ratio. Therefore, our analysis shows how details of the particle shape--beyond their aspect ratio--can be determined by a precise scattering experiment. This is of high relevance in understanding smart materials which involve suspensions of anisotropic colloidal particles.

How does a flexible chain of active particles swell?

We study the swelling of a flexible linear chain composed of active particles by analytical theory and computer simulation. Three different situations are considered: a free chain, a chain confined to an external harmonic trap, and a chain dragged at one end. First, we consider an ideal chain with harmonic springs and no excluded volume between the monomers. The Rouse model of polymers is generalized to the case of self-propelled monomers and solved analytically. The swelling, as characterized by the spatial extension of the chain, scales with the monomer number defining a Flory exponent ν which is ν = 1/2, 0, 1 in the three different situations. As a result, we find that activity does not change the Flory exponent but affects the prefactor of the scaling law. This can be quantitatively understood by mapping the system onto an equilibrium chain with a higher effective temperature such that the chain swells under an increase of the self-propulsion strength. We then use computer simulations to study the effect of self-avoidance on active polymer swelling. In the three different situations, the Flory exponent is now ν = 3/4, 1/4, 1 and again unchanged under self-propulsion. However, the chain extension behaves non-monotonic in the self-propulsion strength.

Transport powered by bacterial turbulence.

We demonstrate that collective turbulentlike motion in a bacterial bath can power and steer the directed transport of mesoscopic carriers through the suspension. In our experiments and simulations, a microwedgelike "bulldozer" draws energy from a bacterial bath of varied density. We obtain that an optimal transport speed is achieved in the turbulent state of the bacterial suspension. This apparent rectification of random motion of bacteria is caused by polar ordered bacteria inside the cusp region of the carrier, which is shielded from the outside turbulent fluctuations.

Circular motion of asymmetric self-propelling particles.

Micron-sized self-propelled (active) particles can be considered as model systems for characterizing more complex biological organisms like swimming bacteria or motile cells. We produce asymmetric microswimmers by soft lithography and study their circular motion on a substrate and near channel boundaries. Our experimental observations are in full agreement with a theory of Brownian dynamics for asymmetric self-propelled particles, which couples their translational and orientational motion.

Dynamics of a deformable active particle under shear flow.

The motion of a deformable active particle in linear shear flow is explored theoretically. Based on symmetry considerations, we propose coupled nonlinear dynamical equations for the particle position, velocity, deformation, and rotation. In our model, both, passive rotations induced by the shear flow as well as active spinning motions, are taken into account. Our equations reduce to known models in the two limits of vanishing shear flow and vanishing particle deformability. For varied shear rate and particle propulsion speed, we solve the equations numerically in two spatial dimensions and obtain a manifold of different dynamical modes including active straight motion, periodic motions, motions on undulated cycloids, winding motions, as well as quasi-periodic and chaotic motions induced at high shear rates. The types of motion are distinguished by different characteristics in the real-space trajectories and in the dynamical behavior of the particle orientation and its deformation. Our predictions can be verified in experiments on self-propelled droplets exposed to a linear shear flow.

Brownian motion of a circle swimmer in a harmonic trap.

We study the dynamics of a Brownian circle swimmer with a time-dependent self-propulsion velocity in an external temporally varying harmonic potential. For several situations, the noise-free swimming paths, the noise-averaged mean trajectories, and the mean-square displacements are calculated analytically or by computer simulation. Based on our results, we discuss optimal swimming strategies in order to explore a maximum spatial range around the trap center. In particular, we find a resonance situation for the maximum escape distance as a function of the various frequencies in the system. Moreover, the influence of the Brownian noise is analyzed by comparing noise-free trajectories at zero temperature with the corresponding noise-averaged trajectories at finite temperature. The latter reveal various complex self-similar spiral or rosette-like patterns. Our predictions can be tested in experiments on artificial and biological microswimmers under dynamical external confinement.

Phototaxis of synthetic microswimmers in optical landscapes.

Many microorganisms, with phytoplankton and zooplankton as prominent examples, display phototactic behaviour, that is, the ability to perform directed motion within a light gradient. Here we experimentally demonstrate that sensing of light gradients can also be achieved in a system of synthetic photo-activated microparticles being exposed to an inhomogeneous laser field. We observe a strong orientational response of the particles because of diffusiophoretic torques, which in combination with an intensity-dependent particle motility eventually leads to phototaxis. Since the aligning torques saturate at high gradients, a strongly rectified particle motion is found even in periodic asymmetric intensity landscapes. Our results are in excellent agreement with numerical simulations of a minimal model and should similarly apply to other particle propulsion mechanisms. Because light fields can be easily adjusted in space and time, this also allows to extend our approach to dynamical environments.

Can the self-propulsion of anisotropic microswimmers be described by using forces and torques?

The self-propulsion of artificial and biological microswimmers (or active colloidal particles) has often been modelled by using a force and a torque entering into the overdamped equations for the Brownian motion of passive particles. This seemingly contradicts the fact that a swimmer is force-free and torque-free, i.e. that the net force and torque on the particle vanish. Using different models for mechanical and diffusiophoretic self-propulsion, we demonstrate here that the equations of motion of microswimmers can be mapped onto those of passive particles with the shape-dependent grand resistance matrix and formally external effective forces and torques. This is consistent with experimental findings on the circular motion of artificial asymmetric microswimmers driven by self-diffusiophoresis. The concept of effective self-propulsion forces and torques significantly facilitates the understanding of the swimming paths, e.g. for a microswimmer under gravity. However, this concept has its limitations when the self-propulsion mechanism of a swimmer is disturbed either by another particle in its close vicinity or by interactions with obstacles, such as a wall.

Gravitaxis of asymmetric self-propelled colloidal particles.

Many motile microorganisms adjust their swimming motion relative to the gravitational field and thus counteract sedimentation to the ground. This gravitactic behaviour 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 which have a homogeneous mass density and a well-defined shape. In experiments and by theoretical modelling, we demonstrate that a shape anisotropy alone is sufficient to induce gravitactic motion with either preferential upward or downward swimming. In addition, also trochoid-like trajectories transversal to the direction of gravity are observed.

Brownian motion and the hydrodynamic friction tensor for colloidal particles of complex shape.

We synthesize colloidal particles with various anisotropic shapes and track their orientationally resolved Brownian trajectories using confocal microscopy. An analysis of appropriate short-time correlation functions provides direct access to the hydrodynamic friction tensor of the particles revealing nontrivial couplings between the translational and rotational degrees of freedom. The results are consistent with calculations of the hydrodynamic friction tensor in the low-Reynolds-number regime for the experimentally determined particle shapes.

Non-Gaussian statistics for the motion of self-propelled Janus particles: experiment versus theory.

Spherical Janus particles are one of the most prominent examples for active Brownian objects. Here, we study the diffusiophoretic motion of such microswimmers in experiment and in theory. Three stages are found: simple Brownian motion at short times, superdiffusion at intermediate times, and finally diffusive behavior again at long times. These three regimes observed in the experiments are compared with a theoretical model for the Langevin dynamics of self-propelled particles with coupled translational and rotational motion. Besides the mean square displacement also higher displacement moments are addressed. In particular, theoretical predictions regarding the non-Gaussian behavior of self-propelled particles are verified in the experiments. Furthermore, the full displacement probability distribution is analyzed, where in agreement with Brownian dynamics simulations either an extremely broadened peak or a pronounced double-peak structure is found, depending on the experimental conditions.

Brownian dynamics of a self-propelled particle in shear flow.

Brownian dynamics of a self-propelled particle in linear shear flow is studied analytically by solving the Langevin equation and in simulation. The particle has a constant propagation speed along a fluctuating orientation and is additionally subjected to a constant torque. In two spatial dimensions, the mean trajectory and the mean square displacement (MSD) are calculated as functions of time t analytically. In general, the mean trajectories are cycloids that are modified by finite temperature effects. With regard to the MSD, different regimes are identified where the MSD scales with t(ν) with ν=0,1,2,3,4. In particular, an accelerated (ν=4) motion emerges if the particle is self-propelled along the gradient direction of the shear flow.