arXiv subjects

Falko Ziebert

Publications and source records attributed to Falko Ziebert.

At least 19 recordsLinked to original sources

arXiv2026

A viscoelastic theory for ultrasound-induced intracellular streaming

Ultrasound is increasingly used to control biological cells, but its multiple physical effects, including radiation force, streaming, heating, and cavitation, require theoretical frameworks that can quantify their relative strengths. Here we develop a semi-analytical model to predict flow patterns and energy distributions of ultrasound-induced intracellular streaming. Cells are modeled as viscoelastic droplets immersed in a fluid with different viscoelastic properties. For this geometry, the momentum equation with an Oldroyd-B constitutive law is solved using a perturbation expansion commonly applied in acoustofluidics. The resulting equations for the different orders of the expansion are solved using partial wave expansion and explicit integration. We investigate our model for a range of parameters which is relevant both for synthetic polymer and for protein solutions. We find a series of flow reversals that we can connect to the distribution of energy into the different modes. We also consider density and compressibility contrasts, which shift these transitions and introduce new ones. Finally, we discuss the potential biological relevance of our mathematical results for cellular sensing and signaling.

physics.bio-ph↗

arXiv2025

A theory for coexistence and selection of branched actin networks in a shared and finite pool of monomers

Cellular actin structures are continuously turned over while keeping similar sizes. Since they all compete for a shared pool of actin monomers, the question arises how they can coexist in these dynamic steady states. Recently, the coexistence of branched actin networks with different densities growing in a shared and finite pool of purified proteins has been demonstrated in a biomimetic bead assay. However, theoretical work in the context of organelle size regulation has mainly been focused on linear architectures, such as single filaments and bundles, and thus is not able to explain this observation. Here we show theoretically that the local depletion of actin monomers caused by the growth of a branched network naturally gives rise to a negative feedback loop between network density and growth rate, and that this competition is captured by one central ordinary differential equation. A comprehensive bifurcation analysis shows that the theory leads to well-defined steady states even in the case of multiple networks sharing the same pool of monomers, without any need for specific molecular processes. Under increasing competition strength, coexistence is replaced by selection. We also show that our theory is in excellent agreement with spatiotemporal simulations, implemented in a finite element framework, and that local depletion even occurs in the presence of a large pool of non-polymerizable actin. In summary, our work suggests that local monomer depletion is the decisive and universal factor controlling growth of branched actin networks.

q-bio.SC↗

arXiv2025

The Hidden Wheel-Within

There is this old, eternal question: Why don't animals have wheels? In this perspective we show that they actually do. And they do so in a physically extraordinary way -- by combining incompatible elasticity, differential geometry and dissipative self-organization. Nature's wheel -- the ``wheel-within'' -- has been mysteriously concealed in plain sight, yet it spins in virtually every slender-body organism: in falling cats, crocodilians spinning to subdue their prey, rolling fruit-fly larvae, circumnutating plants and even in some of our own body movements. Flying somehow under the radar of our cognition, in recent years the wheel-within also tacitly entered the field of soft robotics, finally opening our eyes for its ubiquitous role in Nature. We here identify its underlying physical ingredients, namely the existence of a neutrally-stable, shape-invariant and actively driven elastic mode. We then reflect on various man-made realizations of the wheel-within and outline where it could be spinning from here.

cond-mat.soft↗

arXiv2025

Stochastic Path Integral for the Active Brownian Particle in a Harmonic Potential

In this work we develop and apply a path integral formulation for the microscopic degrees of freedom obeying stochastic differential equations to an active Brownian particle (ABP) trapped in a harmonic potential. The formalism allows to derive exact analytic expressions for the time-dependent moments, like the mean position and the mean square displacement, including full dependence on initial conditions. In addition, the probability distribution of the particle's position can be evaluated systematically as a series expansion in the propulsion speed. Compared to previous methods relying on eigenfunction expansions of the equivalent Fokker-Planck equation, our method is easier to generalize to more complex situations: it does not rely on eigenfunctions but on a reference state that can be solved analytically, which in our case is the passive Brownian particle in a harmonic potential. We exemplify this versatility by also briefly treating an ABP with an active torque (Brownian circle swimmer, BCS) in a harmonic potential.

cond-mat.soft↗

arXiv2025

Active gel theory for cell migration with two myosin isoforms

Myosin II molecular motors slide actin filaments relatively to each other and are essential for force generation, motility and mechanosensing in animal cells. For non-muscle cells, evolution has resulted in three different isoforms, which have different properties concerning the motor cycle and also occur in different abundances in the cells, but their respective biological and physical roles are not fully understood. Here we use active gel theory to demonstrate the complementary roles of isoforms A and B for cell migration. We first show that our model can be derived both from coarse-graining kinetic equations and from nonequilibrium thermodynamics as the macroscopic limit of a two-component Tonks gas. We then parametrize the model and show that motile solutions exist, in which the more abundant and more dynamic isoform A is localized to the front and the stronger isoform B to the rear, in agreement with experiments. Exploring parameter space beyond the isoform parameters typical for animal cells, we also find cell oscillations in length and velocity, which might be realized for genetically engineered systems. We also describe an analytical solution for the stiff limit, which then is used to calculate a state diagram, and the effect of actin polymerization at the boundaries, that leads to an imperfect pitchfork bifurcation. Our findings highlight the importance of including isoform-specific molecular details to describe whole cell behavior.

q-bio.CB↗

arXiv2025

Three-dimensional chiral active Ornstein-Uhlenbeck model for helical motion of microorganisms

Active movement is essential for the survival of microorganisms like bacteria, algae and unicellular parasites. In three dimensions, both swimming and gliding microorganisms often exhibit helical trajectories. One such case are malaria parasites gliding through 3D hydrogels, for which we find that the internal correlation time for the stochastic process generating propulsion is similar to the time taken for one helical turn. Motivated by this experimental finding, here we theoretically analyze the case of finite internal correlation time for microorganisms with helical trajectories as chiral active particles with an Ornstein-Uhlenbeck process for torque. We present an analytical solution which is in very good agreement with computer simulations. We then show that for this type of internal noise, chirality and rotation increase the persistence of motion and results in helical trajectories that have a larger long-time mean squared displacement than straight trajectories at the same propulsion speed. Finally we provide experimental evidence for this prediction for the case of the malaria parasites.

q-bio.CB↗

arXiv2024

The positioning of stress fibers in contractile cells minimizes internal mechanical stress

The mechanics of animal cells is strongly determined by stress fibers, which are contractile filament bundles that form dynamically in response to extracellular cues. Stress fibers allow the cell to adapt its mechanics to environmental conditions and to protect it from structural damage. While the physical description of single stress fibers is well-developed, much less is known about their spatial distribution on the level of whole cells. Here, we combine a finite element method for one-dimensional fibers embedded in an elastic bulk medium with dynamical rules for stress fiber formation based on genetic algorithms. We postulate that their main goal is to achieve minimal mechanical stress in the bulk material with as few fibers as possible. The fiber positions and configurations resulting from this optimization task alone are in good agreement with those found in experiments where cells in 3D-scaffolds were mechanically strained at one attachment point. For optimized configurations, we find that stress fibers typically run through the cell in a diagonal fashion, similar to reinforcement strategies used for composite material.

physics.bio-ph↗

arXiv2024

Active gel model for one-dimensional cell migration coupling actin flow and adhesion dynamics

Migration of animal cells is based on the interplay between actin polymerization at the front, adhesion along the cell-substrate interface, and actomyosin contractility at the back. Active gel theory has been used before to demonstrate that actomyosin contractility is sufficient for polarization and self-sustained cell migration in the absence of external cues, but did not consider the dynamics of adhesion. Likewise, migration models based on the mechanosensitive dynamics of adhesion receptors usually do not include the global dynamics of intracellular flow. Here we show that both aspects can be combined in a minimal active gel model for one-dimensional cell migration with dynamic adhesion. This model demonstrates that load sharing between the adhesion receptors leads to symmetry breaking, with stronger adhesion at the front, and that bistability of migration arises for intermediate adhesiveness. Local variations in adhesiveness are sufficient to switch between sessile and motile states, in qualitative agreement with experiments.

q-bio.CB↗

arXiv2023

Reshaping and Enzymatic Activity may allow Viruses to move through the Mucus

Filamentous viruses like influenza and torovirus often display systematic bends and arcs of mysterious physical origin. We propose that such viruses undergo an instability from a cylindrically symmetric to a toroidally curved state. This ``toro-elastic'' state emerges via spontaneous symmetry breaking under prestress due to short range spike protein interactions magnified by %the filament's surface topography. Once surface stresses are sufficiently large, the filament buckles and the curved state constitutes a soft mode that can potentially propagate through the filament's material frame around a mexican-hat-type potential. In the mucus of our airways, which constitutes a soft, porous 3D network, glycan chains are omnipresent and influenza's spike proteins are known to efficiently bind and cut them. We next show that such a non-equilibrium enzymatic reaction can induce spontaneous rotation of the curved state, leading to a whole body reshaping propulsion similar to -- but different from -- eukaryotic flagella and spirochetes.

cond-mat.soft↗

arXiv2023

Physical phase field model for phagocytosis

We propose and study a simple, physical model for phagocytosis, i.e. the active, actin-mediated uptake of micron-sized particles by biological cells. The cell is described by the phase field method and the driving mechanisms of uptake are actin ratcheting, modeled by a dynamic vector field, as well as cell-particle adhesion due to receptor-ligand binding. We first test the modeling framework for the symmetric situation of a spherical cell engulfing a fixed spherical particle. We then exemplify its versatility by studying various asymmetric situations like different particle shapes and orientations, as well as the simultaneous uptake of two particles. In addition, we perform a perturbation theory of a slightly modified model version in the symmetric setting, allowing to derive a reduced model, shedding light on the effective driving forces and being easier to solve. This work is meant as a first step in describing phagocytosis and we discuss several effects that are amenable to future modeling within the same framework.

cond-mat.soft↗

arXiv2023

The role of the nucleus for cell mechanics: an elastic phase field approach

The nucleus of eukaryotic cells typically makes up around 30% of the cell volume and has significantly different mechanics, which can make it effectively up to ten times stiffer than the surrounding cytoplasm. Therefore it is an important element for cell mechanics, but a quantitative understanding of its mechanical role during whole cell dynamics is largely missing. Here we demonstrate that elastic phase fields can be used to describe dynamical cell processes in adhesive or confining environments in which the nucleus acts as a stiff inclusion. We first introduce and verify our computational method and then study several applications of large relevance. For cells on adhesive patterns, we find that nuclear stress is shielded by the adhesive pattern. For cell compression between two parallel plates, we obtain force-compression curves that allow us to extract an effective modulus for the cell-nucleus composite. For micropipette aspiration, the effect of the nucleus on the effective modulus is found to be much weaker, highlighting the complicated interplay between extracellular geometry and cell mechanics that is captured by our approach. We also show that our phase field approach can be used to investigate the effects of viscoelasticity and cortical tension.

cond-mat.soft↗

arXiv2023

Coarse-graining the vertex model and its response to shear

Tissue dynamics and collective cell motion are crucial biological processes. Their biological machinery is mostly known, and simulation models such as the "active vertex model" (AVM) exist and yield reasonable agreement with experimental observations like tissue fluidization or fingering. However, a good and well-founded continuum description for tissues remains to be developed. In this work we derive a macroscopic description for a two-dimensional cell monolayer by coarse-graining the vertex model through the Poisson bracket approach. We obtain equations for cell density, velocity and the cellular shape tensor. We then study the homogeneous steady states, their stability (which coincides with thermodynamic stability), and especially their behavior under an externally applied shear. Our results contribute to elucidate the interplay between flow and cellular shape. The obtained macroscopic equations present a good starting point for adding cell motion, morphogenetic and other biologically relevant processes.

cond-mat.soft↗

arXiv2022

Optogenetic switching of migration of contractile cells

Cell crawling on flat substrates is based on intracellular flows of the actin cytoskeleton that are driven by both actin polymerization at the front and myosin contractility at the back. The new experimental tool of optogenetics makes it possible to spatially control contraction and thereby possibly also cell migration. Here we analyze this situation theoretically using a one-dimensional active gel model in which the excluded volume interactions of myosin and their aggregation into minifilaments is modeled by a supercritical van der Waals fluid. This physically simple and transparent, but nonlinear and thermodynamically rigorous model predicts bistability between sessile and motile solutions. We then show that one can switch between these two states at realistic parameter ranges via optogenetic activation or inhibition of contractility, in agreement with recent experiments. We also predict the required activation strengths and initiation times.

q-bio.SC↗

arXiv2022

The Physics of Self-Rolling Viruses

Viruses are right at the interface of inanimate matter and life. However, recent experiments [T. Sakai, et al., J.~Virol.~{\bf 92}, e01522-17 (2018)] have shown that some influenza strains can actively roll on glycan-covered surfaces. In a previous letter [F. Ziebert and I. M. Kulić, Phys. Rev. Lett. {\bf 126}, 218101 (2021)] we suggested this to be a form of viral surface metabolism: a collection of spike proteins that attach to and cut the glycans act as a self-organized mechano-chemical motor. Here we study in more depth the physics of the emergent self-rolling states. We give scaling arguments how the motion arises, substantiated by a detailed analytical theory that yields the full torque-angular velocity relation of the self-organized motor. Stochastic Gillespie simulations are used to validate the theory and to quantify stochastic effects like virus detachment and reversals of its direction. Finally, we also cross-check several approximations made previously and show that the proposed mechanism is very robust. All these results point together to the statistical inevitability of viral rolling in presence of enzymatic activity.

cond-mat.soft↗

arXiv2022

Emergent conformational properties of end-tailored transversely propelling polymers

We study the dynamics and conformations of a single active semiflexible polymer whose monomers experience a propulsion force perpendicular to the local tangent, with the end beads being different from the inner beads ("end-tailored"). Using Langevin simulations, we demonstrate that, apart from sideways motion, the relative propulsion strength between the end beads and the polymer backbone significantly changes the conformational properties of the polymers as a function of bending stiffness, end-tailoring and propulsion force. Expectedly, for slower ends the polymer curves away from the moving direction, while faster ends lead to opposite curving, in both cases slightly reducing the center of mass velocity compared to a straight fiber. Interestingly, for faster end beads there is a rich and dynamic morphology diagram: the polymer ends may get folded together to 2D loops or hairpin-like conformations that rotate due to their asymmetry in shape and periodic flapping motion around a rather straight state during full propulsion is also possible. We rationalize the simulations using scaling and kinematic arguments and present the state diagram of the conformations. Sideways propelled fibers comprise a rather unexplored and versatile class of self-propellers, and their study will open novel ways for designing, e.g. motile actuators or mixers in soft robotics.

cond-mat.soft↗

arXiv2021

Optogenetic control of intracellular flows and cell migration: a comprehensive mathematical analysis with a minimal active gel model

The actin cytoskeleton of cells is in continuous motion due to both polymerization of new filaments and their contraction by myosin II molecular motors. Through adhesion to the substrate, such intracellular flow can be converted into cell migration. Recently, optogenetics has emerged as a new powerful experimental method to control both actin polymerization and myosin II contraction. While optogenetic control of polymerization can initiate cell migration by generating protrusion, it is less clear if and how optogenetic control of contraction can also affect cell migration. Here we analyze the latter situation using a minimal variant of active gel theory into which we include optogenetic activation as a spatiotemporally constrained perturbation. The model can describe the symmetrical flow of the actomyosin system observed in optogenetic experiments, but not the long-lasting polarization required for cell migration. Motile solutions become possible if cytoskeletal polymerization is included through the boundary conditions. Optogenetic activation of contraction can then initiate locomotion in a symmetrically spreading cell and strengthen motility in an asymmetrically polymerizing one. If designed appropriately, it can also arrest motility even for protrusive boundaries.

q-bio.SC↗

arXiv2020

How Influenza's Spike Motor Works

While often believed to be a passive agent that merely exploits its host's metabolism, influenza virus has recently been shown to actively move across glycan-coated surfaces. This form of enzymatically driven surface motility is currently not well understood and has been loosely linked to burnt-bridge Brownian ratchet mechanisms. Starting from known properties of influenza's spike proteins, we develop a physical model that quantitatively describes the observed motility. It predicts a collectively emerging dynamics of spike proteins and surface bound ligands that combined with the virus' geometry give rise to a self-organized rolling propulsion. We show that in contrast to a Brownian ratchet, the rotary spike drive is not fluctuation driven but operates optimally as a macroscopic engine in the deterministic regime. The mechanism also applies to relatives of influenza and to man-made analogues like DNA-monowheels and should give guidelines for their optimization.

cond-mat.soft↗

arXiv2020

Dynamics of Fiberboids

Fiberboids are active filaments trapped at the interface of two phases, able of harnessing energy (and matter) fluxes across the interface in order to produce a rolling-like self-propulsion. We discuss several table-top examples and develop the physical framework for understanding their complex dynamics. In spite of some specific features in the examples studied we conclude that the phenomenon of fiberboids is highly generic and robust across different materials, types of fluxes and timescales. Fiberboid motility should play a role from the macroscopic realm down to the micro scale and, as recently hypothesized, possibly as a means of biological self-propulsion that has escaped previous attention.

cond-mat.soft↗