Showing papers by "Frank Jülicher published in 2023"

Liquid spherical shells are a non-equilibrium steady state

Abstract: Liquid-liquid phase separation is the process in which two immiscible liquids demix. This spontaneous phenomenon yields spherical droplets that eventually coarsen to one large, stable droplet governed by the principle of minimal free energy. In chemically fueled phase separation, the formation of phase-separating molecules is coupled to a fuel-driven, nonequilibrium reaction cycle. Chemically fueled phase separation yields dissipative structures sustained by a continuous fuel conversion. Such dissipative structures are ubiquitous in biology but poorly understood as they are governed by non-equilibrium thermodynamics. Here, we bridge the gap between passive, close-to-equilibrium, and active, dissipative structures with chemically fueled phase separation. We observe that spherical, active droplets can transition into a new morphology—a liquid, spherical shell of droplet material. A spherical shell would be highly unstable at equilibrium. Only by continuously converting chemical energy, this dissipative structure can be sustained. We demonstrate the transition mechanism, which is related to the activation of a product outside of the droplet, and the deactivation within the droplets leading to gradients of droplet material. We characterize how far out of equilibrium the spherical shell state is and the chemical power necessary to sustain it. Our work suggests new avenues for assembling complex stable morphologies, which might already be exploited to form membraneless organelles by cells.

Polarized branched Actin modulates cortical mechanics to produce unequal-size daughters during asymmetric division

Abstract: The control of cell shape during cytokinesis requires a precise regulation of mechanical properties of the cell cortex. Only few studies have addressed the mechanisms underlying the robust production of unequal-sized daughters during asymmetric cell division. Here we report that unequal daughter-cell sizes resulting from asymmetric sensory organ precursor divisions in Drosophila are controlled by the relative amount of cortical branched Actin between the two cell poles. We demonstrate this by mistargeting the machinery for branched Actin dynamics using nanobodies and optogenetics. We can thereby engineer the cell shape with temporal precision and thus the daughter-cell size at different stages of cytokinesis. Most strikingly, inverting cortical Actin asymmetry causes an inversion of daughter-cell sizes. Our findings uncover the physical mechanism by which the sensory organ precursor mother cell controls relative daughter-cell size: polarized cortical Actin modulates the cortical bending rigidity to set the cell surface curvature, stabilize the division and ultimately lead to unequal daughter-cell size.

Entrainment of Mutually Synchronized Spatially Distributed 24 GHz Oscillators

Abstract: Synchronization is one of the most challenging aspects of distributed systems in terms of their scalability. Minimal uncertainties can lead to problems or failures regarding data consistency in globally operating data centers or in distributed sensor arrays. Existing approaches to address these challenges are based on hierarchical synchronization concepts which are well understood and have reached technical maturity, but have the disadvantage of having a single point of failure. However, especially for critical infrastructure or backup more resilient solutions are required. Mutual synchronization where oscillators in a network are coupled bidirectionally without a reference have been considered. Due to the flat hierarchy such systems do not have a single point of failure. This work studies how hierarchical synchronization can be combined with architectures implementing mutual synchronization. A network of three mutually coupled 24 GHz oscillators is used to study how injecting a reference signal into one oscillator affects the dynamics. This can be quantified by analyzing in which range of frequencies the network of mutually coupled oscillators can follow the reference frequency. Measurements on a ring and chain network topology forced by an external reference oscillator shown here are in good agreement with the predictions of a nonlinear dynamical model.

Spontaneous flow instabilities of active polar fluids in three dimensions

Abstract: Active polar fluids exhibit spontaneous flow when sufficient active stress is generated by internal molecular mechanisms. This is also referred to as an active Fréedericksz transition. Experiments have revealed the existence of competing in-plane and out-of-plane instabilities in three-dimensional active matter. So far, however, a theoretical model reconciling all observations is missing. In particular, the role of boundary conditions in these instabilities still needs to be explained. Here, we characterize the spontaneous flow transition in a symmetry-preserving three-dimensional active Ericksen-Leslie model, showing that the boundary conditions select the emergent behavior. Using nonlinear numerical solutions and linear perturbation analysis, we explain the mechanism for both in-plane and out-of-plane instabilities under extensile active stress for perpendicular polarity anchoring at the boundary, whereas parallel anchoring only permits in-plane flows under contractile stress or out-of-plane wrinkling under extensile stress.

A cytokinetic ring-driven cell rotation achieves Hertwig’s rule in early development

Abstract: Cells tend to divide along the direction in which they are longest, as famously stated by Oscar Hertwig in 1884 in his ‘long axis’ rule1,2. The orientation of the mitotic spindle determines the cell division axis3, and Hertwig’s long axis rule is usually ensured by forces stemming from microtubules4. Pulling on the spindle from the cell cortex can give rise to unstable behaviors5,6, and we here set out to understand how Hertwig’s long axis rule is realized in early embryonic divisions where cortical pulling forces are prevalent. We focus on early C. elegans development, where we compressed embryos to reveal that cortical pulling forces favor an alignment of the spindle with the cell’s short axis. Strikingly, we find that this misalignment is corrected by an actomyosin-based mechanism that rotates the entire cell, including the mitotic spindle. We uncover that myosin-driven contractility in the cytokinetic ring generates inward forces that align it with the short axis, and thereby the spindle with the long axis. A theoretical model together with experiments using slightly compressed mouse zygotes suggest that a constricting cytokinetic ring can ensure Hertwig’s long axis rule in cells that are free to rotate inside a confining structure, thereby generalizing the underlying principle.

Spontaneous flow instabilities of active polar fluids in three dimensions

Abstract: Active liquid crystals exhibit spontaneous flow when sufficient active stress is generated by internal molecular mechanisms. This is also referred to as active Fr\'{e}edericksz transition. We show this transition in three dimensions and study its dependence on the boundary conditions. Using nonlinear numerical solutions and linear perturbation analysis, we find an out-of-plane transition under extensile active stress for perpendicular polarity anchoring at the boundary, whereas parallel anchoring permits both in-plane flow under contractile stress and out-of-plane wrinkling under extensile stress.