Researcher profile

David Zwicker

David Zwicker contributes to research discovery and scholarly infrastructure.

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Published work

9 published item(s)

preprint2026arXiv

Coexistence of patterned phases in chemically active multicomponent mixtures

Chemically active mixtures exhibit complex patterns that emerge from the interplay of physical interactions and reactions among components. Individually, these two processes are well-understood: Physical interactions can give rise to phase separation, whereas reactions can form reaction-diffusion patterns. To understand the combination of both processes, we identify a Lyapunov functional for a class of chemical reactions. By minimizing this functional, we identify a generalized Gibbs phase rule that governs the number of coexisting patterns, and we demonstrate that complex patterns can be created by the modular combination of independent phases. Our theory unveils complex stationary patterns in chemically active mixtures and provides a framework for analyzing more complex systems.

preprint2026arXiv

Roadmap for Condensates in Cell Biology

Biomolecular condensates govern essential cellular processes yet elude description by traditional equilibrium models. This roadmap, distilled from structured discussions at a workshop and reflecting the consensus of its participants, clarifies key concepts for researchers, funding bodies, and journals. After unifying terminology that often separates disciplines, we outline the core physics of condensate formation, review their biological roles, and identify outstanding challenges in nonequilibrium theory, multiscale simulation, and quantitative in-cell measurements. We close with a forward-looking outlook to guide coordinated efforts toward predictive, experimentally anchored understanding and control of biomolecular condensates.

preprint2022arXiv

Effective simulations of interacting active droplets

Droplets form a cornerstone of the spatiotemporal organization of biomolecules in cells. These droplets are controlled using physical processes like chemical reactions and imposed gradients, which are costly to simulate using traditional approaches, like solving the Cahn-Hilliard equation. To overcome this challenge, we here present an alternative, efficient method. The main idea is to focus on the relevant degrees of freedom, like droplet positions and sizes. We derive dynamical equations for these quantities using analytical solutions to simplified situations. We verify our method against fully-resolved simulations and show that it can describe interacting droplets under the influence of chemical reactions and external gradients using only a fraction of the computational costs of traditional methods. Our method can be extended to include other processes in the future and will thus serve as a relevant platform for understanding the dynamics of droplets in cells.

preprint2022arXiv

Evolved interactions stabilize many coexisting phases in multicomponent liquids

Phase separation has emerged as an essential concept for the spatial organization inside biological cells. However, despite the clear relevance to virtually all physiological functions, we understand surprisingly little about what phases form in a system of many interacting components, like in cells. Here, we introduce a new numerical method based on physical relaxation dynamics to study the coexisting phases in such systems. We use our approach to optimize interactions between components, similar to how evolution might have optimized the interactions of proteins. These evolved interactions robustly lead to a defined number of phases, despite substantial uncertainties in the initial composition, while random or designed interactions perform much worse. Moreover, the optimized interactions are robust to perturbations and they allow fast adaption to new target phase counts. We thus show that genetically encoded interactions of proteins provide versatile control of phase behavior. The phases forming in our system are also a concrete example of a robust emergent property that does not rely on fine-tuning the parameters of individual constituents.

preprint2022arXiv

The intertwined physics of active chemical reactions and phase separation

Phase separation is the thermodynamic process that explains how droplets form in multicomponent fluids. These droplets can provide controlled compartments to localize chemical reactions, and reactions can also affect the droplets' dynamics. This review focuses on the tight interplay between phase separation and chemical reactions originating from thermodynamic constraints. In particular, simple mass action kinetics cannot describe chemical reactions since phase separation requires non-ideal fluids. Instead, thermodynamics implies that passive chemical reactions reduce the complexity of phase diagrams and provide only limited control over the system's behavior. However, driven chemical reactions, which use external energy input to create spatial fluxes, can circumvent thermodynamic constraints. Such active systems can suppress the typical droplet coarsening, control droplet size, and localize droplets. This review provides an extensible framework for describing active chemical reactions in phase separating systems, which forms a basis for improving control in technical applications and understanding self-organized structures in biological cells.

preprint2020arXiv

Elastic stresses reverse Ostwald ripening

When liquid droplets nucleate and grow in a polymer network, compressive stresses can significantly increase their internal pressure, reaching values that far exceed the Laplace pressure. When droplets have grown in a polymer network with a stiffness gradient, droplets in relatively stiff regions of the network tend to dissolve, favoring growth of droplets in softer regions. Here, we show that this elastic ripening can be strong enough to reverse the direction of Ostwald ripening: large droplets can shrink to feed the growth of smaller ones. To numerically model these experiments, we generalize the theory of elastic ripening to account for gradients in solubility alongside gradients in mechanical stiffness.

preprint2020arXiv

Theory of droplet ripening in stiffness gradients

Liquid-liquid phase separation is an important mechanism for compartmentalizing the cell's cytoplasm, allowing the dynamic organization of the components necessary for survival. However, it is not clear how phase separation is affected by the complex viscoelastic environment inside the cell. Here, we study theoretically how stiffness gradients influence droplet growth and arrangement. We show that stiffness gradients imply concentration gradients in the dilute phase, which transport droplet material from stiff to soft regions. Consequently, droplets dissolve in the stiff region, creating a dissolution front. Using a mean-field theory, we predict that the front emerges where the curvature of the elasticity profile is large and that it propagates diffusively. This elastic ripening can occur at much faster rates than classical Ostwald ripening, thus driving the dynamics. Our work shows how gradients in elastic properties control the size and arrangement of droplets, which has potential applications in soft matter physics and plays a role inside biological cells.

preprint2019arXiv

Elastic ripening and inhibition of liquid-liquid phase separation

Phase separation has recently emerged as an important organizational principle in the dense and heterogeneous environment within living cells. Here, we use a synthetic system to show that compressive stresses in a polymer network suppress phase separation of the solvent that swells it. These stresses create a barrier to droplet nucleation that leads to robust, stabilized mixtures well beyond the liquid-liquid phase separation boundary. Network stresses not only alter the stability of mixtures, but they also have a dramatic effect on the ripening of droplets. Gradients in network stresses can drive a new form of ripening, where solute is transported down stiffness gradients. This elastic ripening can be much faster than conventional surface tension driven Ostwald ripening.

preprint2016arXiv

Normalized neural representations of natural odors

The olfactory system removes correlations in natural odors using a network of inhibitory neurons in the olfactory bulb. It has been proposed that this network integrates the response from all olfactory receptors and inhibits them equally. However, how such global inhibition influences the neural representations of odors is unclear. Here, we study a simple statistical model of this situation, which leads to concentration-invariant, sparse representations of the odor composition. We show that the inhibition strength can be tuned to obtain sparse representations that are still useful to discriminate odors that vary in relative concentration, size, and composition. The model reveals two generic consequences of global inhibition: (i) odors with many molecular species are more difficult to discriminate and (ii) receptor arrays with heterogeneous sensitivities perform badly. Our work can thus help to understand how global inhibition shapes normalized odor representations for further processing in the brain.