arXiv subjects

Jens Elgeti

Publications and source records attributed to Jens Elgeti.

At least 19 recordsLinked to original sources

arXiv2025

Free Growth under Tension

Ever since the ground breaking work of Trepat et al. in 2009, we know that cell colonies growing on a substrate can be under tensile mechanical stress. The origin of tension has so far been attributed to cellular motility forces being oriented outward of the colony. Works in the field mainly revolve around how this orientation of the forces can be explained, ranging from velocity alignment, self-sorting due to self-propulsion, to kenotaxis. In this work, we demonstrate that tension in growing colonies can also be explained without cellular motility forces! Using a combination of well established tissue growth simulation technique and analytical modelling, we show how tension can arise as a consequence of simple mechanics of growing tissues. Combining these models with a minimalistic motility model shows how colonies can expand while under even larger tension. Furthermore, our results and analytical models provide novel analysis procedures to identify the underlying mechanics.

physics.bio-ph↗

arXiv2025

The importance of being discrete -- An agent-based model for active nematics and more

We propose an agent-based model of active flexible rods. Inspired by cytoskeletal flows, we introduce activity by an internal flow that contributes to the dissipative forces. The active force between our agents is central and reciprocal, ensuring linear and angular momentum conservation. For nematic activity, we find spontaneous, thresholdless flows and stochastic flow-reorientation, which is accompanied by the formation of topological defects. Defects appear and vanish with activity-dependent rates, and $+1/2$-defects self-propel. These hallmarks of active turbulence are present even on the scale of individual agents. The particle-based interactions lead to an emergent coupling between density and orientation that generates density dipoles around $+1/2$-defects. Finally, we highlight the versatility of our agent-based model by presenting spontaneous flows in three dimensions and tissue growth. Our framework opens the way for an integrated description of living materials, including several forms of activity in the same system.

cond-mat.soft↗

arXiv2024

Viscotaxis of Beating Flagella at Surfaces

Many biological microorganisms and artificial microswimmers react to external cues of environmental gradients by changing their swimming directions. We study here the behavior of eukarytic flagellated microswimmers in linear viscosity gradients. We employ a model of flagellum consisting of a semi-flexible filament with a travelling wave of spontaneous curvature to study viscotaxis of actively beating flagella in two spatial dimensions. The propulsion of the flagellum in a fluid due to a hydrodynamic friction anisotropy is described by resistive-force theory. Using numerical simulations and analytical theory, we show that beating flagella exhibit positive viscotaxis, reorienting themselves toward higher viscosity areas. We quantify this behavior by characterization of the dependence of the rotational velocity on beat amplitude, swimming speed, and wave length. We also examining the effects of asymmetric flagellar wave forms, which imply circular trajectories in the absence of viscosity gradients; here, large asymmetry leads to trochoid-like trajectories perpendicular to the gradient in the form drifting circles. The flagellar deformability strongly reduce the beat amplitude and the viscotatic response. The viscotatic response is shown to be captured by a universal function of the sperm number.

cond-mat.soft↗

arXiv2024

Growth and shrinkage of tissue sheets on substrates: buds, buckles, and pores

Many tissues take the form of thin sheets, being only a single cell thick, but millions of cells wide. These tissue sheets can bend and buckle in the third dimension. In this work, we investigated the growth and shrinkage of suspended and supported tissue sheets using particle-based simulations. We construct a minimum model, combining particle-based tissue growth and meshless membrane models, to simulate the growth of tissue sheets with mechanical feedback. Free suspended growing tissues exhibit wrinkling when growth is sufficiently fast. Conversely, tissues on a substrate form buds when the adhesion to the substrate is weak and/or when the friction with the substrate is strong. These buds undergo a membrane-mediated attraction and subsequently fuse. The complete detachment of tissues from the substrate and straight buckled bump formation are also obtained at very weak adhesion and/or fast growth rates. In the tissue shrinkage, tissue pores grow via Ostwald ripening and coalescence. The reported dynamics can also be applied in research on the detachment dynamics of different tissues with weakened adhesion.

physics.bio-ph↗

arXiv2023

Mechanically-driven Stem Cell Separation in Tissues caused by Proliferating Daughter Cells

The homeostasis of epithelial tissue relies on a balance between the self-renewal of stem cell populations, cellular differentiation, and loss. Although this balance needs to be tightly regulated to avoid pathologies, such as tumor growth, the regulatory mechanisms, both cell-intrinsic and collective, which ensure tissue steady-state are still poorly understood. Here, we develop a computational model that incorporates basic assumptions of stem cell renewal into distinct populations and mechanical interactions between cells. We find that the model generates unexpected dynamic features: stem cells repel each other in the bulk tissue and are thus found rather isolated, as in a number of in vivo contexts. By mapping the system onto a gas of passive Brownian particles with effective repulsive interactions, that arise from the generated flows of differentiated cells, we show that we can quantitatively describe such stem cell distribution in tissues. The interaction potential between a pair of stem cells decays exponentially with a characteristic length that spans several cell sizes, corresponding to the volume of cells generated per stem cell division. Our findings may help understanding the dynamics of normal and cancerous epithelial tissues.

physics.bio-ph↗

arXiv2021

Multi-Ciliated Microswimmers -- Metachronal Coordination and Helical Swimming

The dynamics and motion of multi-ciliated microswimmers with a spherical body and a small number N (with 5 < N < 60) of cilia with length comparable to the body radius, is investigated by mesoscale hydrodynamics simulations. A metachronal wave is imposed for the cilia beat, for which the wave vector has both a longitudinal and a latitudinal component. The dynamics and motion is characterized by the swimming velocity, its variation over the beat cycle, the spinning velocity around the main body axis, as well as the parameters of the helical trajectory. Our simulation results show that the microswimmer motion strongly depends on the latitudinal wave number and the longitudinal phase lag. The microswimmers are found to swim smoothly and usually spin around their own axis. Chirality of the metachronal beat pattern generically generates helical trajectories. In most cases, the helices are thin and stretched, i.e. the helix radius is about an order of magnitude smaller than the pitch. The rotational diffusion of the microswimmer is significantly smaller than the passive rotational diffusion of the body alone, which indicates that the extended cilia contribute strongly to the hydrodynamic radius. The swimming velocity vswim is found to increase with the cilia number N with a slightly sublinear power law, consistent with the behavior expected from the dependence of the transport velocity of planar cilia arrays on the cilia separation.

cond-mat.soft↗

arXiv2020

A minimal model for structure, dynamics, and tension of monolayered cell colonies

The motion of cells in tissues is an ubiquitous phenomenon. In particular, in monolayered cell colonies in vitro, pronounced collective behavior with swirl-like motion has been observed deep within a cell colony, while at the same time, the colony remains cohesive, with not a single cell escaping at the edge. Thus, the colony displays liquid-like properties inside, in coexistence with a cell-free "vacuum" outside. How can adhesion be strong enough to keep cells together, while at the same time not jam the system in a glassy state? What kind of minimal model can describe such a behavior? Which other signatures of activity arise from the internal fluidity? We propose a novel active Brownian particle model with attraction, in which the interaction potential has a broad minimum to give particles enough wiggling space to be collectively in the fluid state. We demonstrate that for moderate propulsion, this model can generate the fluid-vacuum coexistence described above. In addition, the combination of the fluid nature of the colony with cohesion leads to preferred orientation of the cell polarity, pointing outward, at the edge, which in turn gives rise to a tensile stress in the colony -- as observed experimentally for epithelial sheets. For stronger propulsion, collective detachment of cell clusters is predicted. Further addition of an alignment preference of cell polarity and velocity direction results in enhanced coordinated, swirl-like motion, increased tensile stress and cell-cluster detachment.

physics.bio-ph↗

arXiv2020

The role of thickness inhomogeneities in hierarchical cortical folding

For long it has been known that specific patterns of folding are necessary for an optimally functioning brain. For instance, lissencephaly and polymicrogyria can lead to severe mental retardation, short life expectancy, epileptic seizures, and tetraplegia. The construction of a quantitative model on how and why these folds appear is the first step in understanding the cause of these conditions. In recent years, there have been various attempts to understand and model the mechanisms of brain folding. Previous works have shown that mechanical instabilities play a crucial role in the formation of brain folds, and that the geometry of the fetal brain is one of the main factors in dictating the folding characteristics. However, modeling higher-order folding, one of the main characteristics of the human brain, has not been fully tackled. The effects of thickness inhomogeneity in the gyrogenesis of the mammalian brain are studied through finite-element simulations of rectangular slabs. The slabs are divided into two distinct regions, where the outer layer mimics the gray matter, and the inner layer the underlying white matter. Differential growth is introduced by only growing the top layer. The brain tissue is modeled as a neo-Hookean hyperelastic material. Simulations are performed with both, homogeneous and inhomogeneous cortical thickness. The homogeneous cortex is shown to fold into a single wavelength, as common for bilayered materials, while the inhomogeneous cortex folds into more complex conformations: In their early stages of development, structures reminiscent of the deep sulci in the brain are obtained. As the cortex continues to develop, secondary undulations, which are shallower and more variable than the structures obtained in earlier gyrification stage emerge, reproducing well-known characteristics of higher-order folding in the mammalian, and particularly the human, brain.

physics.bio-ph↗

arXiv2020

Instability and fingering of interfaces in growing tissue

Interfaces in tissues are ubiquitous, both between tissue and environment as well as between populations of different cell types. The propagation of an interface can be driven mechanically. % e.g. by a difference in the respective homeostatic stress of the different cell types. Computer simulations of growing tissues are employed to study the stability of the interface between two tissues on a substrate. From a mechanical perspective, the dynamics and stability of this system is controlled mainly by four parameters of the respective tissues: (i) the homeostatic stress (ii) cell motility (iii) tissue viscosity and (iv) substrate friction. For propagation driven by a difference in homeostatic stress, the interface is stable for tissue-specific substrate friction even for very large differences of homeostatic stress; however, it becomes unstable above a critical stress difference when the tissue with the larger homeostatic stress has a higher viscosity. A small difference in directed bulk motility between the two tissues suffices to result in propagation with a stable interface, even for otherwise identical tissues. Larger differences in motility force, however, result in a finite-wavelength instability of the interface. Interestingly, the instability is apparently bound by nonlinear effects and the amplitude of the interface undulations only grows to a finite value in time.

q-bio.TO↗

arXiv2019

Tissue evolution: Mechanical interplay of adhesion, pressure, and heterogeneity

The evolution of various competing cell types in tissues, and the resulting persistent tissue population, is studied numerically and analytically in a particle-based model of active tissues. Mutations change the properties of cells in various ways, including their mechanical properties. Each mutation results in an advantage or disadvantage to grow in the competition between different cell types. While changes in signaling processes and biochemistry play an important role, we focus on changes in the mechanical properties by studying the result of variation of growth force and adhesive cross-interactions between cell types. For independent mutations of growth force and adhesion strength, the tissue evolves towards cell types with high growth force and low internal adhesion strength, as both increase the homeostatic pressure. Motivated by biological evidence, we postulate a coupling between both parameters, such that an increased growth force comes at the cost of a higher internal adhesion strength or vice versa. This tradeoff controls the evolution of the tissue, ranging from unidirectional evolution to very heterogeneous and dynamic populations. The special case of two competing cell types reveals three distinct parameter regimes: Two in which one cell type outcompetes the other, and one in which both cell types coexist in a highly mixed state. Interestingly, a single mutated cell alone suffices to reach the mixed state, while a finite mutation rate affects the results only weakly. Finally, the coupling between changes in growth force and adhesion strength reveals a mechanical explanation for the evolution towards intra-tumor heterogeneity, in which multiple species coexist even under a constant evolutianary pressure.

q-bio.PE↗

arXiv2018

Mechanics of tissue competition: Interfaces stabilize coexistence

Mechanical forces influence the dynamics of growing tissues. Computer simulations are employed to study the importance of interfacial effects in tissue competition. It was speculated that mechanical pressure determines the competition, where the determining quantity is the homeostatic pressure - the pressure where division and apoptosis balance; the tissue with the higher homeostatic pressure overwhelms the other. Surprisingly, a weaker tissue can persist in stable coexistence with a stronger tissue, if adhesion between them is small enough. An analytic continuum description can quantitatively describe the underlying mechanism and reproduce the resulting pressures and cell-number fractions. Computer simulations furthermore display a variety of coexisting structures, ranging from spherical inclusions to a bicontinuous state.

physics.bio-ph↗

arXiv2018

Forces in inhomogeneous open active-particle systems

We study the force that non-interacting point-like active particles apply to a symmetric inert object in the presence of a gradient of activity and particle sources and sinks. We consider two simple patterns of sources and sinks that are common in biological systems. We analytically solve a one dimensional model designed to emulate higher dimensional systems, and study a two dimensional model by numerical simulations. We specify when the particle flux due to the creation and annihilation of particles can act to smooth the density profile that is induced by a gradient in the velocity of the active particles, and find the net resultant force due to both the gradient in activity and the particle flux. These results are compared qualitatively to observations of nuclear motion inside the oocyte, that is driven by a gradient in activity of actin-coated vesicles.

cond-mat.stat-mech↗

arXiv2018

Collective Dynamics of Self-propelled Semiflexible Filaments

The collective behavior of active semiflexible filaments is studied with a model of tangentially driven self-propelled worm-like chains. The combination of excluded-volume interactions and self-propulsion leads to several distinct dynamic phases as a function of bending rigidity, activity, and aspect ratio of individual filaments. We consider first the case of intermediate filament density. For high-aspect-ratio filaments, we identify a transition with increasing propulsion from a state of free-swimming filaments to a state of spiraled filaments with nearly frozen translational motion. For lower aspect ratios, this gas-of-spirals phase is suppressed with growing density due to filament collisions; instead, filaments form clusters similar to self-propelled rods, as activity increases. Finite bending rigidity strongly effects the dynamics and phase behavior. Flexible filaments form small and transient clusters, while stiffer filaments organize into giant clusters, similarly as self-propelled rods, but with a reentrant phase behavior from giant to smaller clusters as activity becomes large enough to bend the filaments. For high filament densities, we identify a nearly frozen jamming state at low activities, a nematic laning state at intermediate activities, and an active-turbulence state at high activities. The latter state is characterized by a power-law decay of the energy spectrum as a function of wave number. The resulting phase diagrams encapsulate tunable non-equilibrium steady states that can be used in the organization of living matter.

physics.bio-ph↗

arXiv2018

Quantitative modelling of nutrient-limited growth of bacterial colonies in microfluidic cultivation

Nutrient gradients and limitations play a pivotal role in the life of all microbes, both in their natural habitat as well as in artificial, microfluidic systems. Spatial concentration gradients of nutrients in densely packed cell configurations may locally affect the bacterial growth leading to heterogeneous micropopulations. A detailed understanding and quantitative modelling of cellular behaviour under nutrient limitations is thus highly desirable. We use microfluidic cultivations to investigate growth and microbial behaviour of the model organism Corynebacterium glutamicum under well-controlled conditions. With a reaction-diffusion type model, parameters are extracted from steady-state experiments with a one-dimensional nutrient gradient. Subsequentially, we employ particle-based simulations with these parameters to predict the dynamical growth of a colony in two dimensions. Comparing the results of those simulations with microfluidic experiments yields excellent agreement. Our modelling approach lays the foundation for a better understanding of dynamic microbial growth processes, both in nature and in applied biotechnology.

physics.bio-ph↗

arXiv2017

Generalized Archimedes' principle in active fluids

We show how a gradient in the motility properties of non-interacting point-like active particles can cause a pressure gradient that pushes a large inert object. We calculate the force on an object inside a system of active particles with position dependent motion parameters, in one and two dimensions, and show that a modified Archimedes' principle is satisfied. We characterize the system, both in terms of the model parameters and in terms of experimentally measurable quantities: the spatial profiles of the density, velocity and pressure. This theoretical analysis is motivated by recent experiments, which showed that the nucleus of a mouse oocyte (immature egg cell) moves from the cortex to the center due to a gradient of activity of vesicles propelled by molecular motors; it more generally applies to artificial systems of controlled localized activity.

cond-mat.stat-mech↗

arXiv2017

Human sperm steer with second harmonics of the flagellar beat

Sperm are propelled by bending waves travelling along the flagellum. During steering in gradients of sensory cues, sperm adjust the flagellar beat waveform. Symmetric and asymmetric beat waveforms produce straight and curved swimming paths, respectively. Two different mechanisms controlling the flagellar beat have been proposed: average intrinsic curvature and dynamic buckling instability. Both mechanisms create spatially asymmetric waveforms that could be modulated for steering. Using video microscopy, we image the flagellar waveform of human sperm tethered with the head to a glass surface. The waveform is characterized by a fundamental beat frequency and its second harmonic. We show that superposition of first and second harmonics breaks the beat symmetry temporally rather than spatially. As a result, sperm rotate around the tethering point. The rotation velocity is determined by the amplitude and phase of the second harmonic. Sperm stimulation with the female sex hormone progesterone enhances the second-harmonic contribution, modulates the flagellar beat, and ultimately sperm rotation. The temporal breaking of beat symmetry represents a new mechanism of sperm steering. Higher-frequency components were also reported for the flagellar beat of other cells; therefore, this steering mechanism might by quite general and could inspire the design of synthetic microswimmers.

physics.bio-ph↗

arXiv2015

Self-propelled Worm-like Filaments: Spontaneous Spiral Formation, Structure, and Dynamics

Worm-like filaments that are propelled homogeneously along their tangent vector are studied by Brownian dynamics simulations. Systems in two dimensions are investigated, corresponding to filaments adsorbed to interfaces or surfaces. A large parameter space covering weak and strong propulsion, as well as flexible and stiff filaments is explored. For strongly propelled and flexible filaments, the free-swimming filaments spontaneously form stable spirals. The propulsion force has a strong impact on dynamic properties, such as the rotational and translational mean square displacement and the rate of conformational sampling. In particular, when the active self-propulsion dominates thermal diffusion, but is too weak for spiral formation, the rotational diffusion coefficient has an activity-induced contribution given by $v_c/ξ_P$, where $v_c$ is the contour velocity and $ξ_P$ the persistence length. In contrast, structural properties are hardly affected by the activity of the system, as long as no spirals form. The model mimics common features of biological systems, such as microtubules and actin filaments on motility assays or slender bacteria, and artificially designed microswimmers.

cond-mat.soft↗

arXiv2015

Giant adsorption of microswimmers: duality of shape asymmetry and wall curvature

The effect of shape asymmetry of microswimmers on their adsorption capacity at confining channel walls is studied by a simple dumbbell model. For a shape polarity of a forward-swimming cone, like the stroke-averaged shape of a sperm, extremely long wall retention times are found, caused by a non-vanishing component of the propulsion force pointing steadily into the wall, which grows exponentially with the self-propulsion velocity and the shape asymmetry. A direct duality relation between shape asymmetry and wall curvature is proposed and verified. Our results are relevant for the design microswimmer with controlled wall-adhesion properties. In addition, we confirm that pressure in active systems is strongly sensitive to the details of the particle-wall interactions.

cond-mat.soft↗