arXiv subjects

Ivar S. Haugerud

Publications and source records attributed to Ivar S. Haugerud.

5 recordsLinked to original sources

arXiv2026

Physical Mechanism of Vacuole Formation in Liquid Droplets

Vacuoles have been observed in liquid droplets across variety of experimental systems, ranging from biomolecular condensates composed of proteins and RNA, to synthetic coacervates formed by charged polymers or synthetic nanostars. These vacuoles are long-lived domains depleted of droplet material, and their formation is puzzling because the associated increase in interfacial area is thermodynamically unfavorable. Using theory, we show that vacuoles form through a generic mechanism: a local spinodal instability within the droplet. We demonstrate this mechanism in several experimentally relevant scenarios, including temperature quenches and droplets coupled to chemical processes occurring either inside or outside the droplet. Using non-equilibrium thermodynamics, we develop a theoretical framework that identifies the physicochemical conditions controlling whether vacuoles form and how big vacuoles can become. Our results suggest molecular designs and chemical pathways that promote vacuolation, enabling multi-compartment formation with engineered functions such as enhanced surface catalysis and compartment fission.

cond-mat.soft↗

arXiv2026

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.

physics.bio-ph↗

arXiv2025

Harvesting chemical power from cyclic environments

Life relies on a sophisticated metabolic molecular machinery that turns over high-energy molecules to evolve complex macromolecules and assemblies. At the molecular origin of life, such machinery was absent, implying the need for simple yet robust physical mechanisms to harvest energy from the environment and perform chemical work or produce chemical power. However, the mechanisms involved in harvesting energy from a macroscopic cyclic environment to drive chemical processes on the molecular scale remain elusive. In this work, we propose a theory that describes the kinetics of chemical reactions in a system subject to a cyclic reservoir with varying properties. We compare cycles of solvent (wet-dry cycles), with cycles of a component participating in a chemical reaction (reactant cycle). We find that for both wet-dry and reactant cycles, resonance frequencies exist at which the chemical power is maximal. We identify which cycle type is more beneficial in harvesting chemical power for different molecular interactions. Our findings of harvest efficiencies around ten percent suggest that the cyclic environment could have played a key role in catalyzing the metabolic molecular machinery at the molecular origin of life.

physics.chem-ph↗

arXiv2025

Excitability and oscillations of active droplets

In living cells, cycles of formation and dissolution of liquid droplets can mediate biological functions such as DNA repair. However, the minimal physicochemical prerequisite for such droplet oscillations remains elusive. Here, we present a simple model composed of only two independent chemical components with their diffusive and chemical fluxes governed by non-equilibrium thermodynamics. There is turnover of fuel that maintains a chemical reaction away from equilibrium, leading to active droplets. We find that a single active droplet undergoes a pitchfork-bifurcation in the droplet volume upon increasing the fueling strength. Strikingly, the active droplet becomes excitable upon adding a further chemical reaction. For sufficient fueling, the system undergoes self-sustained oscillations cycling between droplet formation and dissolution. The minimal nature of our model suggests self-sustained active droplets as functional modules for de novo life.

cond-mat.soft↗

arXiv2024

Theory for sequence selection via phase separation and oligomerization

Non-equilibrium selection pressures were proposed for the formation of oligonucleotides with rich functionalities encoded in their sequences, such as catalysis. Since phase separation was shown to direct various chemical processes, we ask whether condensed phases can provide mechanisms for sequence selection. To answer this question, we use non-equilibrium thermodynamics and describe the reversible oligomerization of different monomers to sequences at non-dilute conditions prone to phase separation. We find that when sequences oligomerize, their interactions give rise to phase separation, boosting specific sequences' enrichment and depletion. Our key result is that phase separation gives rise to a selection pressure for the oligomerization of specific sequence patterns when fragmentation maintains the system away from equilibrium. Specifically, slow fragmentation favors alternating sequences that interact well with their environment (more cooperative), while fast fragmentation selects sequences with extended motifs capable of specific sequence interactions (less cooperative). Our results highlight that out-of-equilibrium condensed phases could provide versatile hubs for Darwinian-like evolution toward functional sequences, both relevant for the molecular origin of life and de novo life.

cond-mat.stat-mech↗