arXiv subjects

Erwin Frey

Publications and source records attributed to Erwin Frey.

At least 37 records · Page 2Linked to original sources

arXiv2024

Availability versus carrying capacity: Phases of asymmetric exclusion processes competing for finite pools of resources

We address how the interplay between the finite availability and carrying capacity of particles at different parts of a spatially extended system can control the steady state currents and density profiles in the one-dimensional current-carrying lanes connecting the different parts of the system. To study this, we set up a minimal model consisting of two particle reservoirs of the same finite carrying capacity connected by two equally sized anti-parallel asymmetric exclusion processes (TASEP). We focus on the steady-state currents and particle density profiles in the two TASEP lanes. The ensuing phases and the phase diagrams, which can be remarkably complex, are parametrized by the model parameters defining particle exchange between the TASEP lanes and the reservoirs and the filling fraction of the particles that determine the total resources available. These parameters may be tuned to make the densities of the two TASEP lanes globally uniform or piece-wise continuous in the form of a combination of a single localized domain wall and a spatially constant density or a pair of delocalized domain walls. Our model reveals that the two reservoirs can be preferentially populated or depopulated in the steady states.

cond-mat.stat-mech↗

arXiv2024

Dimensionality reduction in bulk-boundary reaction-diffusion systems

Intracellular protein patterns regulate many vital cellular functions, such as the processing of spatiotemporal information or the control of shape deformations. To do so, pattern-forming systems can be sensitive to the cell geometry by means of coupling the protein dynamics on the cell membrane to dynamics in the cytosol. Recent studies demonstrated that modeling the cytosolic dynamics in terms of an averaged protein pool disregards possibly crucial aspects of the pattern formation, most importantly concentration gradients normal to the membrane. At the same time, the coupling of two domains (surface and volume) with different dimensions renders many standard tools for the numerical analysis of self-organizing systems inefficient. Here, we present a generic framework for projecting the cytosolic dynamics onto the lower-dimensional surface that respects the influence of cytosolic concentration gradients in static and evolving geometries. This method uses a priori physical information about the system to approximate the cytosolic dynamics by a small number of dominant characteristic concentration profiles (basis), akin to basis transformations of finite element methods. As a proof of concept, we apply our framework to a toy model for volume-dependent interrupted coarsening, evaluate the accuracy of the results for various basis choices, and discuss the optimal basis choice for biologically relevant systems. Our analysis presents an efficient yet accurate method for analysing pattern formation with surface-volume coupling in evolving geometries.

physics.bio-ph↗

arXiv2024

Design principles for fast and efficient self-assembly processes

Self-assembly is a fundamental concept in biology and of significant interest to nanotechnology. Significant progress has been made in characterizing and controlling the properties of the resulting structures, both experimentally and theoretically. However, much less is known about kinetic constraints and determinants of dynamical properties like time efficiency, although these constraints can become severe limiting factors of self-assembly processes. Here we investigate how the time efficiency and other dynamical properties of reversible self-assembly depend on the morphology (shape) of the building blocks for systems in which the binding energy between the constituents is large. As paradigmatic examples, we stochastically simulate the self-assembly of constituents with triangular, square, and hexagonal morphology into two-dimensional structures of a specified size. We find that the constituents' morphology critically determines the assembly time and how it scales with the size of the target structure. Our analysis reveals three key structural parameters defined by the morphology: The nucleation size and attachment order, which describe the effective order of the chemical reactions by which clusters nucleate and grow, respectively, and the growth exponent, which determines how the growth rate of an emerging structure scales with its size. Using this characterization, we formulate an effective theory of the self-assembly kinetics, which we show exhibits an inherent scale invariance. This allows us to identify general scaling laws that describe the minimal assembly time as a function of the size of the target structure. We show how these insights on the kinetics of self-assembly processes can be used to design assembly schemes that could significantly increase the time efficiency and robustness of artificial self-assembly processes.

cond-mat.soft↗

arXiv2024

The role of mobility in epidemics near criticality

The general epidemic process (GEP), also known as susceptible-infected-recovered model (SIR), describes how an epidemic spreads within a population of susceptible individuals who acquire permanent immunization upon recovery. This model exhibits a second-order absorbing state phase transition, commonly studied assuming immobile healthy individuals. We investigate the impact of mobility on disease spreading near the extinction threshold by introducing two generalizations of GEP, where the mobility of susceptible and recovered individuals is examined independently. In both cases, including mobility violates GEP's rapidity reversal symmetry and alters the number of absorbing states. The critical dynamics of the models are analyzed through a perturbative renormalization group approach and large-scale stochastic simulations using a Gillespie algorithm. The renormalization group analysis predicts both models to belong to the same novel universality class describing the critical dynamics of epidemic spreading when the infected individuals interact with a diffusive species and gain immunization upon recovery. At the associated renormalization group fixed point, the immobile species decouples from the dynamics of the infected species, dominated by the coupling with the diffusive species. Numerical simulations in two dimensions affirm our renormalization group results by identifying the same set of critical exponents for both models. Violation of the rapidity reversal symmetry is confirmed by breaking the associated hyperscaling relation. Our study underscores the significance of mobility in shaping population spreading dynamics near the extinction threshold.

cond-mat.stat-mech↗

arXiv2024

Coarsening and wavelength selection far from equilibrium: a unifying framework based on singular perturbation theory

Intracellular protein patterns are described by (nearly) mass-conserving reaction-diffusion systems. While these patterns initially form out of a homogeneous steady state due to the well-understood Turing instability, no general theory exists for the dynamics of fully nonlinear patterns. We develop a unifying theory for wavelength-selection dynamics in (nearly) mass-conserving two-component reaction-diffusion systems independent of the specific mathematical model chosen. This encompasses both the dynamics of the mesa- and peak-shaped patterns found in these systems. Our analysis uncovers a diffusion- and a reaction-limited regime of the dynamics, which provides a systematic link between the dynamics of mass-conserving reaction-diffusion systems and the Cahn-Hilliard as well as conserved Allen-Cahn equations, respectively. A stability threshold in the family of stationary patterns with different wavelengths predicts the wavelength selected for the final stationary pattern. At short wavelengths, self-amplifying mass transport between single pattern domains drives coarsening while at large wavelengths weak source terms that break strict mass conservation lead to an arrest of the coarsening process. The rate of mass competition between pattern domains is calculated analytically using singular perturbation theory, and rationalized in terms of the underlying physical processes. The resulting closed-form analytical expressions enable us to quantitatively predict the coarsening dynamics and the final pattern wavelength. We find excellent agreement of these expressions with numerical results. The systematic understanding of the length-scale dynamics of fully nonlinear patterns in two-component systems provided here builds the basis to reveal the mechanisms underlying wavelength selection in multi-component systems with potentially several conservation laws.

nlin.PS↗

arXiv2024

Supramolecular assemblies in active motor-filament systems: micelles, bilayers, and foams

Active matter systems evade the constraints of thermal equilibrium, leading to the emergence of intriguing collective behavior. A paradigmatic example is given by motor-filament mixtures, where the motion of motor proteins drives alignment and sliding interactions between filaments and their self-organization into macroscopic structures. After defining a microscopic model for these systems, we derive continuum equations, exhibiting the formation of active supramolecular assemblies such as micelles, bilayers and foams. The transition between these structures is driven by a branching instability, which destabilizes the orientational order within the micelles, leading to the growth of bilayers at high microtubule densities. Additionally, we identify a fingering instability, modulating the shape of the micelle interface at high motor densities. We study the role of various mechanisms in these two instabilities, such as contractility, active splay, and anchoring, allowing for generalization beyond the system considered here.

cond-mat.soft↗

arXiv2023

Anomalous Collective Dynamics of Auto-Chemotactic Populations

While the role of local interactions in nonequilibrium phase transitions is well studied, a fundamental understanding of the effects of long-range interactions is lacking. We study the critical dynamics of reproducing agents subject to autochemotactic interactions and limited resources. A renormalization group analysis reveals distinct scaling regimes for fast (attractive or repulsive) interactions; for slow signal transduction, the dynamics is dominated by a diffusive fixed point. Furthermore, we present a correction to the Keller-Segel nonlinearity emerging close to the extinction threshold and a novel nonlinear mechanism that stabilizes the continuous transition against the emergence of a characteristic length scale due to a chemotactic collapse.

cond-mat.stat-mech↗

arXiv2023

Hierarchical defect-induced condensation in active nematics

Topological defects play a central role in the formation and organization of various biological systems. Historically, such nonequilibrium defects have been mainly studied in the context of homogeneous active nematics. Phase-separated systems, in turn, are known to form dense and dynamic nematic bands, but typically lack topological defects. In this paper, we use agent-based simulations of weakly aligning, self-propelled polymers and demonstrate that contrary to the existing paradigm phase-separated active nematics form ${-1/2}$ defects. Moreover, these defects, emerging due to interactions among dense nematic bands, constitute a novel second-order collective state. We investigate the morphology of defects in detail and find that their cores correspond to a strong increase in density, associated with a condensation of nematic fluxes. Unlike their analogs in homogeneous systems, such condensed defects form and decay in a different way and do not involve positively charged partners. We additionally observe and characterize lateral arc-like structures that separate from a band's bulk and move in transverse direction. We show that the key control parameters defining the route from stable bands to the coexistence of dynamic lanes and defects are the total density of particles and their path persistence length. We introduce a hydrodynamic theory that qualitatively recapitulates all the main features of the agent-based model, and use it to show that the emergence of both defects and arcs can be attributed to the same anisotropic active fluxes. Finally, we present a way to artificially engineer and position defects, and speculate about experimental verification of the provided model.

cond-mat.soft↗

arXiv2023

Periodic temporal environmental variations induce coexistence in resource competition models

Natural ecosystems, in particular on the microbial scale, are inhabited by a large number of species. The population size of each species is affected by interactions of individuals with each other and by spatial and temporal changes in environmental conditions, such as resource abundance. Here, we use a generic population dynamics model to study how, and under what conditions, a periodic temporal environmental variation can alter an ecosystem's composition and biodiversity. We demonstrate that using time scale separation allows one to qualitatively predict the long-term population dynamics of interacting species in varying environments. We show that the notion of Tilman's R* rule, a well-known principle that applies for constant environments, can be extended to periodically varying environments if the time scale of environmental changes (e.g., seasonal variations) is much faster than the time scale of population growth (doubling time in bacteria). When these time scales are similar, our analysis shows that a varying environment deters the system from reaching a steady state, and stable coexistence between multiple species becomes possible. Our results posit that biodiversity can in part be attributed to natural environmental variations.

q-bio.PE↗

arXiv2023

Enzyme-enriched condensates show self-propulsion, positioning, and coexistence

Enzyme-enriched condensates can organize the spatial distribution of their substrates by catalyzing non-equilibrium reactions. Conversely, an inhomogeneous substrate distribution induces enzyme fluxes through substrate-enzyme interactions. We find that condensates move towards the center of a confining domain when this feedback is weak. Above a feedback threshold, they exhibit self-propulsion, leading to oscillatory dynamics. Moreover, catalysis-driven enzyme fluxes can lead to interrupted coarsening, resulting in equidistant condensate positioning, and to condensate division.

physics.bio-ph↗

arXiv2022

Bridging scales in a multiscale pattern-forming system

Self-organized pattern formation is vital for many biological processes. Reaction-diffusion models have advanced our understanding of how biological systems develop spatial structures, starting from homogeneity. However, biological processes inherently involve multiple spatial and temporal scales and transition from one pattern to another over time, rather than progressing from homogeneity to a pattern. To deal with such multiscale systems, coarse-graining methods are needed that allow the dynamics to be reduced to the relevant degrees of freedom at large scales, but without losing information about the patterns at the small scales. Here, we present a semi-phenomenological approach which exploits mass-conservation in pattern formation, and enables to reconstruct information about patterns from the large-scale dynamics. The basic idea is to partition the domain into distinct regions (coarse-grain) and determine instantaneous dispersion relations in each region, which ultimately inform about local pattern-forming instabilities. We illustrate our approach by studying the Min system, a paradigmatic model for protein pattern formation. By performing simulations, we first show that the Min system produces multiscale patterns in a spatially heterogeneous geometry. This prediction is confirmed experimentally by in vitro reconstitution of the Min system. Using a recently developed theoretical framework for mass-conserving reaction-diffusion systems, we show that the spatiotemporal evolution of the total protein densities on large-scales reliably predicts the pattern-forming dynamics. Our approach provides an alternative and versatile theoretical framework for complex systems where analytical coarse-graining methods are not applicable, and can in principle be applied to a wide range of systems with an underlying conservation law.

physics.bio-ph↗

arXiv2022

Geometry-induced patterns through mechanochemical coupling

Intracellular protein patterns regulate a variety of vital cellular processes such as cell division and motility, which often involve dynamic changes of cell shape. These changes in cell shape may in turn affect the dynamics of pattern-forming proteins, hence leading to an intricate feedback loop between cell shape and chemical dynamics. While several computational studies have examined the resulting rich dynamics, the underlying mechanisms are not yet fully understood. To elucidate some of these mechanisms, we explore a conceptual model for cell polarity on a dynamic one-dimensional manifold. Using concepts from differential geometry, we derive the equations governing mass-conserving reaction-diffusion systems on time-evolving manifolds. Analyzing these equations mathematically, we show that dynamic shape changes of the membrane can induce pattern-forming instabilities in parts of the membrane, which we refer to as regional instabilities. Deformations of the local membrane geometry can also (regionally) suppress pattern formation and spatially shift already existing patterns. We explain our findings by applying and generalizing the local equilibria theory of mass-conserving reaction-diffusion systems. This allows us to determine a simple onset criterion for geometry-induced pattern-forming instabilities, which is linked to the phase-space structure of the reaction-diffusion system. The feedback loop between membrane shape deformations and reaction-diffusion dynamics then leads to a surprisingly rich phenomenology of patterns, including oscillations, traveling waves, and standing waves that do not occur in systems with a fixed membrane shape. Our work reveals that the local conformation of the membrane geometry acts as an important dynamical control parameter for pattern formation in mass-conserving reaction-diffusion systems.

nlin.PS↗

arXiv2022

Control of protein-based pattern formation via guiding cues

Proteins control many vital functions in living cells, such as cell growth and cell division. Reliable coordination of these functions requires the spatial and temporal organizaton of proteins inside cells, which encodes information about the cell's geometry and the cell-cycle stage. Such protein patterns arise from protein transport and reaction kinetics, and they can be controlled by various guiding cues within the cell. Here, we review how protein patterns are guided by cell size and shape, by other protein patterns that act as templates, and by the mechanical properties of the cell. The basic mechanisms of guided pattern formation are elucidated with reference to recent observations in various biological model organisms. We posit that understanding the controlled formation of protein patterns in cells will be an essential part of understanding information processing in living systems.

physics.bio-ph↗

arXiv2021

Length Regulation Drives Self-Organization in Filament-Motor Mixtures

Cytoskeletal networks form complex intracellular structures. Here we investigate a minimal model for filament-motor mixtures in which motors act as depolymerases and thereby regulate filament length. Combining agent-based simulations and hydrodynamic equations, we show that resource-limited length regulation drives the formation of filament clusters despite the absence of mechanical interactions between filaments. Even though the orientation of individual remains fixed, collective filament orientation emerges in the clusters, aligned orthogonal to their interfaces.

physics.bio-ph↗

arXiv2021

Subdiffusive Activity Spreading in the Diffusive Epidemic Process

The diffusive epidemic process is a paradigmatic example of an absorbing state phase transition in which healthy and infected individuals spread with different diffusion constants. Using stochastic activity spreading simulations in combination with finite-size scaling analyses we reveal two qualitatively different processes that characterize the critical dynamics: subdiffusive propagation of infection clusters and diffusive fluctuations in the healthy population. This suggests the presence of a strong-coupling regime and sheds new light on a longstanding debate about the theoretical classification of the system.

cond-mat.stat-mech↗

arXiv2021

Snowdrift game induces pattern formation in systems of self-propelled particles

Evolutionary games between species are known to lead to intriguing spatio-temporal patterns in systems of diffusing agent. However, the role of inter-species interactions is hardly studied when agents are (self-)propelled, as is the case in many biological systems. Here, we combine aspects from active-matter and evolutionary game theory and study a system of two species whose individuals are (self-)propelled and interact through a snowdrift game. We derive hydrodynamic equations for the density and velocity fields of both species from which we identify parameter regimes in which one or both species form macroscopic orientational order as well as regimes of propagating wave patterns. Interestingly, we find simultaneous wave patterns in both species that result from the interplay between alignment and snowdrift interactions - a feedback mechanism that we call game-induced pattern formation. We test these results in agent-based simulations and confirm the different regimes of order and spatio-temporal patterns as well as game-induced pattern formation.

physics.bio-ph↗

arXiv2021

Wavelength selection by interrupted coarsening in reaction-diffusion systems

Wavelength selection in reaction--diffusion systems can be understood as a coarsening process that is interrupted by counteracting processes at certain wavelengths. We first show that coarsening in mass-conserving systems is driven by self-amplifying mass transport between neighboring high-density domains. We derive a general coarsening criterion and show that coarsening is generically uninterrupted in two-component systems that conserve mass. The theory is then generalized to study interrupted coarsening and anti-coarsening due to weakly-broken mass conservation, providing a general path to analyze wavelength selection in pattern formation far from equilibrium.

nlin.PS↗

arXiv2021

Diffusive coupling of two well-mixed compartments elucidates elementary principles of protein-based pattern formation

Spatial organization of proteins in cells is important for many biological functions. In general, the nonlinear, spatially coupled models for protein-pattern formation are only accessible to numerical simulations, which has limited insight into the general underlying principles. To overcome this limitation, we adopt the setting of two diffusively coupled, well-mixed compartments that represents the elementary feature of any pattern -- an interface. For intracellular systems, the total numbers of proteins are conserved on the relevant timescale of pattern formation. Thus, the essential dynamics is the redistribution of the globally conserved mass densities between the two compartments. We present a phase-portrait analysis in the phase-space of the redistributed masses that provides insights on the physical mechanisms underlying pattern formation. We demonstrate this approach for several paradigmatic model systems. In particular, we show that the pole-to-pole Min oscillations in Escherichia coli are relaxation oscillations of the MinD polarity orientation. This reveals a close relation between cell polarity oscillatory patterns in cells. Critically, our findings suggest that the design principles of intracellular pattern formation are found in characteristic features in these phase portraits (nullclines and fixed points). These features are not uniquely determined by the topology of the protein-interaction network but depend on parameters (kinetic rates, diffusion constants) and distinct networks can give rise to equivalent phase portrait features.

nlin.PS↗