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Oliver Bäumchen

Publications and source records attributed to Oliver Bäumchen.

At least 19 recordsLinked to original sources

Probing molecular-motor forces driving Chlamydomonas gliding motility by holographic microscopy

The unicellular microalga Chlamydomonas reinhardtii is a widely used model microswimmer for biophysicists. However, the interest around C. reinhardtii goes beyond its swimming capabilities. In fact, light can drastically alter its behavior: under blue illumination, the cell attaches to a nearby surface and intermittently glides on it. Such a gliding motility is powered by molecular-motor proteins operating in the cell's cilia, and the related machinery has established the cell as a prime reference for the study of intraciliary-transport mechanisms. In the present work, we track label-free cells using in-line holographic microscopy, which allows us to characterize their gliding motility without requiring fluorescent labeling. After benchmarking the experimental technique on swimming cells, we show that, while gliding, the cells move mostly along the surface, and exhibit anomalous-diffusive features, including Lorentzian-like distributions of displacements. Furthermore, gliding trajectories, by being intermittent, are relevant candidates to determine forces at the molecular-motor scale. We report a gliding threshold of about 20~pN, which is compatible with the activity of single molecular motors, and on the same order of magnitude as the forces generated during intraciliary cargo transport.

cond-mat.soft↗

Metabolic activity controls the emergence of coherent flows in microbial suspensions

Photosynthetic microbes have evolved and successfully adapted to the ever-changing environmental conditions in complex microhabitats throughout almost all ecosystems on Earth. In the absence of light, they can sustain their biological functionalities through aerobic respiration, and even in anoxic conditions through anaerobic metabolic activity. For a suspension of photosynthetic microbes in an anaerobic environment, individual cellular motility is directly controlled by its photosynthetic activity, i.e. the intensity of the incident light absorbed by chlorophyll. The effects of the metabolic activity on the collective motility on the population level, however, remain elusive so far. Here, we demonstrate that at high light intensities, a suspension of photosynthetically active microbes exhibits a stable reverse sedimentation profile of the cell density due to the microbes' natural bias to move against gravity. With decreasing photosynthetic activity, and therefore suppressed individual motility, the living suspension becomes unstable giving rise to coherent bioconvective flows. The collective motility is fully reversible and manifests as regular, three-dimensional plume structures, in which flow rates and cell distributions are directly controlled via the light intensity. The coherent flows emerge in the highly unfavourable condition of lacking both light and oxygen and, thus, might help the microbial collective to expand the exploration of their natural habitat in search for better survival conditions.

physics.bio-ph↗

Quantifying gliding forces of filamentous cyanobacteria by self-buckling

Filamentous cyanobacteria are one of the oldest and today still most abundant lifeforms on earth, with manifold implications in ecology and economics. Their flexible filaments, often several hundred cells long, exhibit gliding motility in contact with solid surfaces. The underlying force generating mechanism is not yet understood. Here, we demonstrate that propulsion forces and friction coefficients are strongly coupled in the gliding motility of filamentous cyanobacteria. We directly measure their bending moduli using micropipette force sensors, and quantify propulsion and friction forces by analyzing their self-buckling behavior, complemented with analytical theory and simulations. The results indicate that slime extrusion unlikely generates the gliding forces, but support adhesion-based hypotheses, similar to the better-studied single-celled myxobacteria. The critical self-buckling lengths align well with the peaks of natural length distributions, indicating the importance of self-buckling for the organization of their collective in natural and artificial settings.

physics.bio-ph↗

Emergent probability fluxes in confined microbial navigation

When the motion of a motile cell is observed closely, it appears erratic, and yet the combination of nonequilibrium forces and surfaces can produce striking examples of organization in microbial systems. While most of our current understanding is based on bulk systems or idealized geometries, it remains elusive how and at which length scale self-organization emerges in complex geometries. Here, using experiments, analytical and numerical calculations we study the motion of motile cells under controlled microfluidic conditions, and demonstrate that probability flux loops organize active motion even at the level of a single cell exploring an isolated compartment of nontrivial geometry. By accounting for the interplay of activity and interfacial forces, we find that the boundary's curvature determines the nonequilibrium probability fluxes of the motion. We theoretically predict a universal relation between fluxes and global geometric properties that is directly confirmed by experiments. Our findings open the possibility to decipher the most probable trajectories of motile cells and may enable the design of geometries guiding their time-averaged motion.

cond-mat.soft↗

Light-regulated adsorption and desorption of Chlamydomonas cells at surfaces

Microbial colonization of surfaces represents the first step towards biofilm formation, which is a recurring phenomenon in nature with beneficial and detrimental implications in technological and medical settings. Consequently, there is a current interest in elucidating the fundamental aspects of the initial stages of biofilm formation of microorganisms on solid surfaces. While most of the research is oriented to understand bacterial surface colonization, such observations at a fundamental level using photosynthetic microalgae are thus far elusive. Recent single-cell studies showed that the flagellar adhesion of Chlamydomonas is switched on in blue light and switched off under red light [Kreis et al., Nature Physics, 2018, 14, 45-49]. Here, we study this light-switchable surface association of C. reinhardtii on the population level and measure the kinetics of adsorption and desorption of suspensions of motile cells on glass surfaces using bright field optical microscopy. We observe that both processes exhibit a response lag relative to the time at which the blue- and red-light conditions are set and model this feature using time-delayed Langmuir-type kinetics. We find that cell adsorption occurs significantly faster than desorption, which we attribute to the protein-mediated molecular adhesion mechanism of the cells. Adsorption experiments using phototactically blind Chlamydomonas mutants demonstrate that phototaxis does not affect the cell adsorption kinetics. Hence, this method can be used as an assay for characterizing the dynamics of the surface colonization of microbial species exhibiting light-regulated surface adhesion.

physics.bio-ph↗

Motility and self-organization of gliding Chlamydomonas populations

Cellular appendages such as cilia and flagella represent universal tools enabling cells and microbes, among other essential functionalities, to propel themselves in diverse environments. In its planktonic, i.e. freely swimming, state the unicellular bi-flagellated microbe Chlamydomonas reinhardtii employs a periodic breaststroke-like flagellar beating to displace the surrounding fluid. Another flagella-mediated motility mode is observed for surface-associated Chlamydomonas cells, which glide along the surface by means of force transduction through an intraflagellar transport machinery. Experiments and statistical motility analysis demonstrate that this gliding motility enhances clustering and supports self-organization of Chlamydomonas populations. We employ Minkowski functionals to characterize the spatiotemporal organization of the surface-associated cell monolayer. We find that simulations based on a purely mechanistic approach cannot capture the observed non-random cell configurations. Quantitative agreement with experimental data however is achieved when considering a minimal cognitive model of the flagellar mechanosensing.

physics.bio-ph↗

Measuring and upscaling micromechanical interactions in a cohesive granular material

The mechanical properties of a disordered heterogeneous medium depend, in general, on a complex interplay between multiple length scales. Connecting local interactions to macroscopic observables, such as stiffness or fracture, is thus challenging in this type of material. Here, we study the properties of a cohesive granular material composed of glass beads held together by soft polymer bridges. We characterise the mechanical response of single bridges under traction and shear, using a setup based on the deflection of flexible micropipettes. These measurements, along with information from X-ray microtomograms of the granular packings, then inform large-scale discrete element model (DEM) simulations. Although simple, these simulations are constrained in every way by empirical measurement and accurately predict mechanical responses of the aggregates, including details on their compressive failure, and how the material's stiffness depends on the stiffness and geometry of its parts. By demonstrating how to accurately relate microscopic information to macroscopic properties, these results provide new perspectives for predicting the behaviour of complex disordered materials, such as porous rock, snow, or foam.

cond-mat.soft↗

Light controls motility and phase separation of photosynthetic microbes

Large ensembles of interacting, out-of-equilibrium agents are a paradigm of active matter. Their constituents' intrinsic activity may entail the spontaneous separation into localized phases of high and low densities. Motile microbes, equipped with ATP-fueled engines, are prime examples of such phase-separating active matter, which is fundamental in myriad biological processes. The fact that spontaneous spatial aggregation is not widely recognized as a general feature of microbial communities challenges the generalisation of phase separation beyond artificial active systems. Here, we report on the phase separation of populations of Chlamydomonas reinhardtii that can be controlled by light in a fully reversible manner. We trace this phenomenon back to the light- and density-dependent motility, thus bridging the gap from light perception on the single-cell level to collective spatial self-organization into regions of high and low density. Its spectral sensitivity suggests that microbial motility and phase separation are regulated by the activity of the photosynthetic machinery. Characteristic fingerprints of the stability and dynamics of this active system paint a picture that cannot be reconciled with the current physical understanding of phase separation in artificial active matter, whereby collective behavior can emerge from inherent motility modulation in response to changing stimuli. Our results therefore point towards the existence of a broader class of self-organization phenomena in living systems.

physics.bio-ph↗

Dynamic Force Measurements on Swimming Chlamydomonas Cells using Micropipette Force Sensors

Flagella and cilia are cellular appendages that inherit essential functions of microbial life including sensing and navigating the environment. In order to propel a swimming microorganism they displace the surrounding fluid by means of periodic motions, while precisely-timed modulations of their beating patterns enable the cell to steer towards or away from specific locations. Characterizing the dynamic forces, however, is challenging and typically relies on indirect experimental approaches. Here, we present direct in vivo measurements of the dynamic forces of motile Chlamydomonas reinhardtii cells in controlled environments. The experiments are based on partially aspirating a living microorganism at the tip of a micropipette force sensor and optically recording the micropipette's position fluctuations with high temporal and sub-pixel spatial resolution. We provide an analytic elasto-hydrodynamic model for the micropipette force sensor and describe how to obtain the micropipette's full frequency response function from a dynamic force calibration. Using this approach, we find dynamic forces during the free swimming activity of individual Chlamydomonas reinhardtii cells of 23$\pm$5 pN resulting from the coordinated flagellar beating with a frequency of 51$\pm$6 Hz. In addition to measurements in bulk liquid environment, we study the dynamic forces of the biflagellated microswimmer in the vicinity of a solid/liquid interface. As we gradually decrease the distance of the swimming microbe to the interface, we measure a significantly enhanced force transduction at distances larger than the maximum extend of the beating flagella, highlighting the importance of hydrodynamic interactions for scenarios in which flagellated microorganisms encounter surfaces.

physics.bio-ph↗

Adsorption-Induced Slip Inhibition for Polymer Melts on Ideal Substrates

Hydrodynamic slip of a liquid at a solid surface represents a fundamental phenomenon in fluid dynamics that governs liquid transport at small scales. For polymeric liquids, de Gennes predicted that the Navier boundary condition together with the theory of polymer dynamics imply extraordinarily large interfacial slip for entangled polymer melts on ideal surfaces; this Navier-de Gennes model was confirmed using dewetting experiments on ultra-smooth, low-energy substrates. Here, we use capillary leveling - surface tension driven flow of films with initially non-uniform thickness - of polymeric films on these same substrates. Measurement of the slip length from a robust one-parameter fit to a lubrication model is achieved. We show that at the lower shear rates involved in leveling experiments as compared to dewetting ones, the employed substrates can no longer be considered ideal. The data is instead consistent with physical adsorption of polymer chains at the solid/liquid interface. We extend the Navier-de Gennes description using one additional parameter, namely the density of physically adsorbed chains per unit surface. The resulting formulation is found to be in excellent agreement with the experimental observations.

cond-mat.soft↗

Elastocapillary Levelling of Thin Viscous Films on Soft Substrates

A thin liquid film with non-zero curvature at its free surface spontaneously flows to reach a flat configuration, a process driven by Laplace pressure gradients and resisted by the liquid's viscosity. Inspired by recent progresses on the dynamics of liquid droplets on soft substrates, we here study the relaxation of a viscous film supported by an elastic foundation. Experiments involve thin polymer films on elastomeric substrates, where the dynamics of the liquid-air interface is monitored using atomic force microscopy. A theoretical model that describes the coupled evolution of the solid-liquid and the liquid-air interfaces is also provided. In this soft-levelling configuration, Laplace pressure gradients not only drive the flow, but they also induce elastic deformations on the substrate that affect the flow and the shape of the liquid-air interface itself. This process represents an original example of elastocapillarity that is not mediated by the presence of a contact line. We discuss the impact of the elastic contribution on the levelling dynamics and show the departure from the classical self-similarities and power laws observed for capillary levelling on rigid substrates.

cond-mat.soft↗

Nucleated dewetting in supported ultra-thin liquid films with hydrodynamic slip

This study reveals the influence of the surface energy and solid/liquid boundary condition on the breakup mechanism of dewetting ultra-thin polymer films. Using silane self-assembled monolayers, SiO$_2$ substrates are rendered hydrophobic and provide a strong slip rather than a no-slip solid/liquid boundary condition. On undergoing these changes, the thin-film breakup morphology changes dramatically -- from a spinodal mechanism to a breakup which is governed by nucleation and growth. The experiments reveal a dependence of the hole density on film thickness and temperature. The combination of lowered surface energy and hydrodynamic slip brings the studied system closer to the conditions encountered in bursting unsupported films. As for unsupported polymer films, a critical nucleus size is inferred from a free energy model. This critical nucleus size is supported by the film breakup observed in the experiments using high speed \emph{in situ} atomic force microscopy.

cond-mat.soft↗

Curvature-guided motility of microalgae in geometric confinement

Microorganisms, such as bacteria and microalgae, often live in habitats consisting of a liquid phase and a plethora of interfaces. The precise ways in which these motile microbes behave in their confined environment remain unclear. Using experiments, Brownian dynamics simulations, and analytical theory, we study the motility of a single Chlamydomonas microalga in an isolated microhabitat with controlled geometric properties. We demonstrate how the geometry of the habitat controls the cell's navigation in confinement. The probability of finding the cell swimming near the boundary scales linearly with the wall curvature, as seen for both circular and elliptical chambers. The theory, utilizing an asymmetric dumbbell model of the cell and steric wall interactions, captures this curvature-guided navigation quantitatively with no free parameters.

physics.bio-ph↗

Universal contact-line dynamics at the nanoscale

The relaxation dynamics of the contact angle between a viscous liquid and a smooth substrate is studied at the nanoscale. Through atomic force microscopy measurements of polystyrene nanostripes we monitor simultaneously the temporal evolution of the liquid-air interface as well as the position of the contact line. The initial configuration exhibits high curvature gradients and a non-equilibrium contact angle that drive liquid flow. Both these conditions are relaxed to achieve the final state, leading to three successive regimes along time: i) stationary-contact-line levelling; ii) receding-contact-line dewetting; iii) collapse of the two fronts. For the first regime, we reveal the existence of a self-similar evolution of the liquid interface, which is in excellent agreement with numerical calculations from a lubrication model. For different liquid viscosities and film thicknesses we provide evidence for a transition to dewetting featuring a universal critical contact angle and dimensionless time.

cond-mat.soft↗

Relaxation and Intermediate Asymptotics of a Rectangular Trench in a Viscous Film

The surface of a thin liquid film with nonconstant curvature flattens as a result of capillary forces. While this leveling is driven by local curvature gradients, the global boundary conditions greatly influence the dynamics. Here, we study the evolution of rectangular trenches in a polystyrene nanofilm. Initially, when the two sides of a trench are well separated, the asymmetric boundary condition given by the step height controls the dynamics. In this case, the evolution results from the leveling of two noninteracting steps. As the steps broaden further and start to interact, the global symmetric boundary condition alters the leveling dynamics. We report on full agreement between theory and experiments for: the capillary-driven flow and resulting time dependent height profiles; a crossover in the power-law dependence of the viscous energy dissipation as a function of time as the trench evolution transitions from two noninteracting to interacting steps; and the convergence of the profiles to a universal self-similar attractor that is given by the Green's function of the linear operator describing the dimensionless linearized thin film equation.

cond-mat.soft↗

Numerical solutions of thin film equations for polymer flows

We report on the numerical implementation of thin film equations that describe the capillary-driven evolution of viscous films, in two-dimensional configurations. After recalling the general forms and features of these equations, we focus on two particular cases inspired by experiments: the leveling of a step at the free surface of a polymer film, and the leveling of a polymer droplet over an identical film. In each case, we first discuss the long-term self-similar regime reached by the numerical solution before comparing it to the experimental profile. The agreement between theory and experiment is excellent, thus providing a versatile probe for nanorheology of viscous liquids in thin film geometries.

cond-mat.soft↗

Capillary leveling of stepped films with inhomogeneous molecular mobility

A homogeneous thin polymer film with a stepped height profile levels due to the presence of Laplace pressure gradients. Here we report on studies of polymeric samples with precisely controlled, spatially inhomogeneous molecular weight distributions. The viscosity of a polymer melt strongly depends on the chain length distribution; thus, we learn about thin-film hydrodynamics with viscosity gradients. These gradients are achieved by stacking two films with different molecular weights atop one another. After a sufficient time these samples can be well described as having one dimensional viscosity gradients in the plane of the film, with a uniform viscosity normal to the film. We develop a hydrodynamic model that accurately predicts the shape of the experimentally observed self-similar profiles. The model allows for the extraction of a capillary velocity, the ratio of the surface tension and the viscosity, in the system. The results are in excellent agreement with capillary velocity measurements of uniform mono- and bi-disperse stepped films and are consistent with bulk polymer rheology.

cond-mat.soft↗

Solid Surface Structure Affects Liquid Order at the Polystyrene/SAM Interface

We present a combined x-ray and neutron reflectivity study characterizing the interface between polystyrene (PS) and silanized surfaces. Motivated by the large difference in slip velocity of PS on top of dodecyl-trichlorosilane (DTS) and octadecyl-trichlorosilane (OTS) found in previous studies, these two systems were chosen for the present investigation. The results reveal the molecular conformation of PS on silanized silicon. Differences in the molecular tilt of OTS and DTS are replicated by the adjacent phenyl rings of the PS. We discuss our findings in terms of a potential link between the microscopic interfacial structure and dynamic properties of polymeric liquids at interfaces.

cond-mat.soft↗