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Joakim Stenhammar

Publications and source records attributed to Joakim Stenhammar.

At least 19 recordsLinked to original sources

Pseudo-giant number fluctuations and nematic order in microswimmer suspensions

Giant number fluctuations (GNFs), whereby the standard deviation $ΔN$ in the local number of particles $\langle N \rangle$ grows faster than $\sqrt{\langle N \rangle}$, are a hallmark property of dry active matter systems with orientational order, such as a collection of granular particles on a vibrated plate. This contrasts with momentum-conserving ("wet") active matter systems, such as suspensions of swimming bacteria, where no theoretical prediction of GNFs exist, although numerous experimental observations of such enhanced fluctuations have been reported. In this Letter, we numerically confirm the emergence of super-Gaussian number fluctuations in a 3-dimensional suspension of pusher microswimmers undergoing a transition to collective motion. These fluctuations emerge sharply above the transition, but only for sufficiently large values of the bacterial persistence length $\ell_p = v_s / λ$, where $v_s$ is the bacterial swimming speed and $λ$ the tumbling rate. Crucially, these "pseudo-GNFs" differ from true GNFs, as they only occur on length scales shorter than the typical size $ξ$ of nematic patches in the collective motion state, which is in turn proportional to the single-swimmer persistence length $\ell_p$. Our results thus suggest that observations of enhanced density fluctuations in biological active matter systems actually represent transient effects that decay away beyond mesoscopic length scales, and raises the question to what extent "true" GNFs with universal properties can exist in the presence of fluid flows.

cond-mat.soft

Collective motion in a sheet of microswimmers

Self-propelled micron-size particles suspended in a fluid, like bacteria or synthetic microswimmers, are strongly non-equilibrium systems where particle motility breaks the microscopic detailed balance, often resulting in large-scale collective motion. Previous theoretical work has identified long-range hydrodynamic interactions as the main driver of collective motion in unbounded dilute suspension of rear-actuated ("pusher") microswimmers. In contrast, most experimental studies of collective motion in microswimmer suspensions have been carried out in quasi-2-dimensional geometries such as in thin films or near solid or fluid interfaces, where both the swimmers' motion and their long-range flow fields become altered due to the proximity of a boundary. Here, we study numerically a minimal model of microswimmers in such a restricted geometry, where the particles move in the midplane between two no-slip walls. For pushers, we demonstrate collective motion with only short-ranged order, in contrast with the long-ranged flows observed in unbounded systems. For front-actuated ("puller") microswimmers, we discover a long-wavelength density instability resulting in the formation of dense microswimmer clusters. Both types of collective motion are fundamentally different from their previously studied counterparts in unbounded domains. Our results illustrate that hydrodynamic screening due to the presence of a wall is subdominant in determining the collective state of the suspension, which is instead dictated by the geometrical restriction of the swimmers' motion.

cond-mat.soft

Hydrodynamic instabilities in a 2-D sheet of microswimmers embedded in a 3-D fluid

A collection of microswimmers immersed in an incompressible fluid is characterised by strong interactions due to the long-range nature of the hydrodynamic fields generated by individual organisms. As a result, suspensions of rear-actuated `pusher' swimmers such as bacteria exhibit a collective motion state often referred to as `bacterial turbulence', characterised by large-scale chaotic flows. The onset of collective motion in pusher suspensions is classically understood within the framework of mean-field kinetic theories for dipolar swimmers. In bulk 2-D and 3-D, the theory predicts that the instability leading to bacterial turbulence is due to mutual swimmer reorientation and sets in at the largest length scale available to the suspension. Here, we construct a similar kinetic theory for the case of a dipolar microswimmer suspension restricted to a two-dimensional plane embedded in a three-dimensional incompressible fluid. This setting qualitatively mimics the effect of swimming close to a two-dimensional interface. We show that the in-plane flow fields are effectively compressible in spite of the incompressibility of the 3-D bulk fluid, and that microswimmers on average act as sources (pushers) or sinks (pullers). We analyse stability of the homogeneous and isotropic state, and find two types of instability that are qualitatively different from the bulk, three-dimensional case: First, we show that the analogue of the orientational pusher instability leading to bacterial turbulence in bulk systems instead occurs at the smallest length-scale available to the system. Second, an instability associated with density variations arises in puller suspensions as a generic consequence of the effective in-plane compressibility. We conclude that confinement can have a crucial role in determining the collective behaviour of microswimmer suspensions.

cond-mat.soft

Interplay between Brownian and hydrodynamic tracer diffusion in suspensions of swimming microorganisms

The general problem of tracer diffusion in non-equilibrium baths is important in a wide range of systems, from the cellular level to geographical lengthscales. In this paper, we revisit the archetypical example of such a system: a collection of small passive particles immersed in a dilute suspension of non-interacting dipolar microswimmers, representing bacteria or algae. In particular, we consider the interplay between thermal (Brownian) diffusion and hydrodynamic (active) diffusion due to the persistent advection of tracers by microswimmer flow fields. Previously, it has been argued that even a moderate amount of Brownian diffusion is sufficient to significantly reduce the persistence time of tracer advection, leading to a significantly reduced value of the effective active diffusion coefficient $D_A$ compared to the non-Brownian case. Here, we show by large-scale simulations and kinetic theory that this effect is in fact only practically relevant for microswimmers that effectively remain stationary while still stirring up the surrounding fluid, so-called \emph{shakers}. In contrast, for moderate and high values of the swimming speed $v_s$, relevant for biological microswimmer suspensions, the effect of Brownian motion on $D_A$ is negligible, leading to the effects of advection by microswimmers and Brownian motion being additive. This conclusion contrasts with previous results from the literature, and encourages a reinterpretation of recent experimental measurements of $D_A$ for tracer particles of varying size in bacterial suspensions.

cond-mat.soft

Pair-distribution function of active Brownian spheres in three spatial dimensions: simulation results and analytical representation

The pair-distribution function, which provides information about correlations in a system of interacting particles, is one of the key objects of theoretical soft matter physics. In particular, it allows for microscopic insights into the phase behavior of active particles. While this function is by now well studied for two-dimensional active matter systems, the more complex and more realistic case of three-dimensional systems is not well understood by now. In this work, we analyze the full pair-distribution function of spherical active Brownian particles interacting via a Weeks-Chandler-Andersen potential in three spatial dimensions using Brownian dynamics simulations. Besides extracting the structure of the pair-distribution function from the simulations, we obtain an analytical representation for this function, parametrized by activity and concentration, which takes into account the symmetries of a homogeneous stationary state. Our results are useful as input to quantitative models of active Brownian particles and advance our understanding of the microstructure in dense active fluids.

cond-mat.soft

Thermal fluctuations and osmotic stability of lipid vesicles

Biological membranes constantly change their shape in response to external stimuli, and understanding the remodeling and stability of vesicles in heterogeneous environments is therefore of fundamental importance for a range of cellular processes. One crucial question is how vesicles respond to external osmotic stresses, imposed by differences in solute concentrations between the vesicle interior and exterior. Previous analyses of the membrane bending energy have predicted that micron-sized giant unilamellar vesicles (GUVs) should become globally deformed already for nanomolar concentration differences, in contrast to experimental findings that find deformations at much higher osmotic stresses. In this article, we analyze the mechanical stability of a spherical vesicle exposed to an external osmotic pressure in a statistical-mechanical model, including the effect of thermally excited membrane bending modes. We find that the inclusion of thermal fluctuations of the vesicle shape changes renders the vesicle deformation continuous, in contrast to the abrupt transition in the athermal picture. Crucially, however, the predicted critical pressure associated with global vesicle deformation remains the same as when thermal fluctuations are neglected, approximately six orders of magnitude smaller than the typical collapse pressure recently observed experimentally for GUVs. We conclude by discussing possible sources of this persisting dissonance between theory and experiments.

cond-mat.soft

Anisotropic diffusion of ellipsoidal tracers in microswimmer suspensions

Tracer particles immersed in suspensions of biological microswimmers such as E. coli or C. reinhardtii display phenomena unseen in conventional equilibrium systems, including strongly enhanced diffusivity relative to the Brownian value and non-Gaussian displacement statistics. In dilute, 3-dimensional suspensions, these phenomena have typically been explained by the hydrodynamic advection of point tracers by isolated microswimmers, while, at higher concentrations, correlations between pusher microswimmers such as E. coli can increase the effective diffusivity even further. Anisotropic tracers in active suspensions can be expected to exhibit even more complex behaviour than spherical ones, due to the presence of a nontrivial translation-rotation coupling. Using large-scale lattice Boltzmann simulations of model microswimmers described by extended force dipoles, we study the motion of ellipsoidal point tracers immersed in 3-dimensional microswimmer suspensions. We find that the rotational diffusivity of tracers is much less affected by swimmer-swimmer correlations than the translational diffusivity. We furthermore study the anisotropic translational diffusion in the particle frame and find that, in pusher suspensions, the diffusivity along the ellipsoid major axis is higher than in the direction perpendicular to it, albeit with a smaller ratio than for Brownian diffusion. Thus, we find that far field hydrodynamics cannot account for the anomalous coupling between translation and rotation observed in experiments, as was recently proposed. Finally, we study the probability distributions (PDFs) of translational and rotational displacements. In accordance with experimental observations, for short observation times we observe strongly non-Gaussian PDFs that collapse when rescaled with their variance, which we attribute to the ballistic nature of tracer motion at short times.

cond-mat.soft

Collective guiding of acoustically propelled nano- and microparticles for medical applications

One of the most important potential applications of self-propelled nano- and microdevices is targeted drug delivery. To realize this, biocompatible particles that can be guided collectively towards a target inside a patient's body are required. Acoustically propelled nano- and microparticles constitute a promising candidate for such biocompatible, artificial motile particles. The main remaining obstacle to targeted drug delivery by self-propelled nano- and microdevices is to also achieve a reliable and biocompatible method for guiding them collectively to their target. Here, we propose such a method. As we confirm by computer simulations, it allows for the remote guiding of large numbers of acoustically propelled particles to a prescribed target by combining a space- and time-dependent acoustic field and a time-dependent magnetic field. The method works without detailed knowledge about the particle positions and for arbitrary initial particle distributions. With these features, it paves the way for the future application of self-propelled particles as vehicles for targeted drug delivery in nanomedicine.

cond-mat.soft

An Introduction to Motility-Induced Phase Separation

Motility-induced phase separation, MIPS, is arguably the most well-studied collective phenomenon occurring in active matter without alignment interactions (scalar active matter). Its basic origin is simple: since self-propelled particles accumulate where they move slowly, having a propulsion speed that decreases steeply enough with density, due to collisions or chemical interactions, leads to a feedback loop that induces the formation of a dense phase. In these notes, I will discuss some of the main theoretical and computational efforts that have been made over the last decade in understanding the basic structural and dynamical properties of MIPS phase coexistence in microscopic active particle models.

cond-mat.soft

ArGSLab: A tool for analyzing experimental or simulated particle networks

Microscopy and particle-based simulations are both powerful techniques to study aggregated particulate matter such as colloidal gels. The data provided by these techniques often contains information on a wide array of length scales, but structural analysis methods typically focus on the local particle arrangement, even though the data also contains information about the particle network on the mesoscopic length scale. In this paper, we present a MATLAB software package for quantifying mesoscopic network structures in colloidal samples. ArGSLab (Arrested and Gelated Structures Laboratory) extracts a network backbone from the input data, which is in turn transformed into a set of nodes and links for graph theory-based analysis. The routines can process both image stacks from microscopy as well as explicit coordinate data, and thus allows quantitative comparison between simulations and experiments. ArGSLab furthermore enables the accurate analysis of microscopy data where, e.g., an extended point spread function prohibits the resolution of individual particles. We demonstrate the resulting output for example datasets from both microscopy and simulation of colloidal gels, in order to showcase the capability of ArGSLab to quantitatively analyze data from various sources. The freely available software package can be used either with a provided graphical user interface or directly as a MATLAB script.

cond-mat.soft

Crystal-to-Crystal Transitions in Binary Mixtures of Soft Colloids

In this article, we demonstrate a method for inducing reversible crystal-to-crystal transitions in binary mixtures of soft colloidal particles. Through a controlled decrease of salinity and increasingly dominating electrostatic interactions, a single sample is shown to reversibly organize into entropic crystals, electrostatic attraction-dominated crystals or aggregated gels, which we quantify using microscopy and image analysis. We furthermore analyze crystalline structures with bond order analysis to discern between two crystal phases. We observe the different phases using a sample holder geometry that allows both in situ salinity control and imaging through Confocal Laser Scanning Microscopy, and apply a synthesis method producing particles with high resolvability in microscopy with control over particle size. The particle softness provides for an enhanced crystallization speed, while altering the re-entrant melting behavior as compared to hard sphere systems. This work thus provides several tools for use in the reproducible manufacture and analysis of binary colloidal crystals.

cond-mat.soft

Using patchy particles to prevent local rearrangements in models of non-equilibrium colloidal gels

Simple models based on isotropic interparticle attractions often fail to capture experimentally observed structures of colloidal gels formed through spinodal decomposition and subsequent arrest: the resulting gels are typically denser and less branched than their experimental counterparts. Here we simulate gels formed from soft particles with directional attractions ("patchy particles"), designed to inhibit lateral particle rearrangement after aggregation. We directly compare simulated structures with experimental colloidal gels made using soft attractive microgel particles, by employing a "skeletonization" method that reconstructs the 3-dimensional backbone from experiment or simulation. We show that including directional attractions with sufficient valency leads to strongly branched structures compared to isotropic models. Furthermore, combining isotropic and directional attractions provides additional control over aggregation kinetics and gel structure. Our results show that the inhibition of lateral particle rearrangements strongly affects the gel topology, and is an important effect to consider in computational models of colloidal gels.

cond-mat.soft

Swimming suppresses correlations in dilute suspensions of pusher microorganisms

Active matter exhibits various forms of non-equilibrium states in the absence of external forcing, including macroscopic steady-state currents. Such states are often too complex to be modelled from first principles and our understanding of their physics relies heavily on minimal models. These have mostly been studied in the case of "dry" active matter, where particle dynamics are dominated by friction with their surroundings. Significantly less is known about systems with long-range hydrodynamic interactions that belong to "wet" active matter. Dilute suspensions of motile bacteria, modelled as self-propelled dipolar particles interacting solely through long-ranged hydrodynamic fields, are arguably the most studied example from this class of active systems. Their phenomenology is well-established: at sufficiently high density of bacteria, there appear large-scale vortices and jets comprising many individual organisms, forming a chaotic state commonly known as bacterial turbulence. As revealed by computer simulations, below the onset of collective motion, the suspension exhibits very strong correlations between individual microswimmers stemming from the long-ranged nature of dipolar fields. Here we demonstrate that this phenomenology is captured by the minimal model of microswimmers. We develop a kinetic theory that goes beyond the commonly used mean-field assumption, and explicitly takes into account such correlations. Notably, these can be computed exactly within our theory. We calculate the fluid velocity variance, spatial and temporal correlation functions, the fluid velocity spectrum, and the enhanced diffusivity of tracer particles. We find that correlations are suppressed by particle self-propulsion, although the mean-field behaviour is not restored even in the limit of very fast swimming.

cond-mat.soft

Symmetric mixtures of pusher and puller microswimmers behave as noninteracting suspensions

Suspensions of rear- and front-actuated microswimmers immersed in a fluid, known respectively as ``pushers'' and ``pullers'', display qualitatively different collective behaviours: beyond a characteristic density, pusher suspensions exhibit a hydrodynamic instability leading to collective motion known as active turbulence, a phenomenon which is absent for pullers. In this Letter, we describe the collective dynamics of a binary pusher--puller mixture using kinetic theory and large-scale particle-resolved simulations. We derive and verify an instability criterion, showing that the critical density for active turbulence moves to higher values as the fraction $χ$ of pullers is increased and disappears for $χ\geq 0.5$. We then show analytically and numerically that the two-point hydrodynamic correlations of the 1:1 mixture are equal to those of a suspension of noninteracting swimmers. Strikingly, our numerical analysis furthermore shows that the full probability distribution of the fluid velocity fluctuations collapses onto the one of a noninteracting system at the same density, where swimmer--swimmer correlations are strictly absent. Our results thus indicate that the fluid velocity fluctuations in 1:1 pusher--puller mixtures are exactly equal to those of the corresponding noninteracting suspension at any density, a surprising cancellation with no counterpart in equilibrium long-range interacting systems.

cond-mat.soft

Anisotropic dynamics of a self-assembled colloidal chain in an active bath

Anisotropic macromolecules exposed to non-equilibrium (active) noise are very common in biological systems, and an accurate understanding of their anisotropic dynamics is therefore crucial. Here, we experimentally investigate the dynamics of isolated chains assembled from magnetic microparticles at a liquid-air interface and moving in an active bath consisting of motile E. coli bacteria. We investigate both the internal chain dynamics and the anisotropic center-of-mass dynamics through particle tracking. We find that both the internal and center-of-mass dynamics are greatly enhanced compared to the passive case, i.e., a system without bacteria, and that the center-of-mass diffusion coefficient $D$ features a non-monotonic dependence as a function of the chain length. Furthermore, our results show that the relationship between the components of $D$ parallel and perpendicular with respect to the direction of the applied magnetic field is preserved in the active bath compared to the passive case, with a higher diffusion in the parallel direction, in contrast to previous findings in the literature. We argue that this qualitative difference is due to subtle differences in the experimental geometry and conditions and the relative roles played by long-range hydrodynamic interactions and short-range collisions.

cond-mat.soft

Pair-distribution function of active Brownian spheres in two spatial dimensions: simulation results and analytic representation

We investigate the full pair-distribution function of a homogeneous suspension of spherical active Brownian particles interacting by a Weeks-Chandler-Andersen potential in two spatial dimensions. The full pair-distribution function depends on three coordinates describing the relative positions and orientations of two particles, the Péclet number specifying the activity of the particles, and their mean packing density. This five-dimensional function is obtained from Brownian dynamics simulations. We discuss its structure taking into account all of its degrees of freedom. In addition, we present an approximate analytic expression for the product of the full pair-distribution function and the interparticle force. We find that the analytic expression, which is typically needed when deriving analytic models for the collective dynamics of active Brownian particles, is in good agreement with the simulation results. The results of this work can thus be expected to be helpful for the further theoretical investigation of active Brownian particles as well as nonequilibrium statistical physics in general.

cond-mat.soft

Derivation of the Derjaguin approximation for the case of inhomogeneous solvents

The Derjaguin approximation (DA) relates the force between curved surfaces to the interaction free energy between parallel planes. It is typically derived by considering the direct interaction between the bodies involved, thus treating the effect of an intervening solvent implicitly by a rescaling of the corresponding Hamaker constant. Here, we provide a generalization of DA to the case of a molecular medium between the bodies, as is the case in most applications. The derivation is based on an explicit statistical-mechanical treatment of the contribution to the interaction force from a molecular solvent using a general expression for intermolecular and molecule-surface interactions. Starting from an exact expression for the force, DA is arrived at by a series of well-defined approximations. Our results show that DA remains valid in a molecular solvent as long as (i) the surface-molecule interactions are of much shorter range than the radius R of the sphere and (ii) the density correlation length in the solvent is smaller than R. We then extend our analysis to the case where a phase transition occurs between the surfaces, which cannot easily be covered using a statistical-mechanical formalism due to the discontinuous change in the density of the medium. Instead using a continuum thermodynamic description, we show that this phase transformation induces an attractive force between the bodies, and that the force between curved surfaces can be related to the free energy in the corresponding planar case, in accordance with DA.

cond-mat.soft

Particle-resolved lattice Boltzmann simulations of 3-dimensional active turbulence

Collective behaviour in suspensions of microswimmers is often dominated by the impact of long-ranged hydrodynamic interactions. These phenomena include active turbulence, where suspensions of pusher bacteria at sufficient densities exhibit large-scale, chaotic flows. To study this collective phenomenon, we use large-scale (up to $N=3\times 10^6$) particle-resolved lattice Boltzmann simulations of model microswimmers described by extended stresslets. Such system sizes enable us to obtain quantitative information about both the transition to active turbulence and characteristic features of the turbulent state itself. In the dilute limit, we test analytical predictions for a number of static and dynamic properties against our simulation results. For higher swimmer densities, where swimmer-swimmer interactions become significant, we numerically show that the length- and timescales of the turbulent flows increase steeply near the predicted finite-system transition density.

cond-mat.soft