Role of hydrodynamic flows in chemically driven droplet division

We study the hydrodynamics and shape changes of chemically active droplets. In non-spherical droplets, surface tension generates hydrodynamic flows that drive liquid droplets into a spherical shape. Here we show that spherical droplets that are maintained away from thermodynamic equilibrium by chemical reactions may not remain spherical but can undergo a shape instability which can lead to spontaneous droplet division. In this case chemical activity acts against surface tension and tension-induced hydrodynamic flows. By combining low Reynolds-number hydrodynamics with phase separation dynamics and chemical reaction kinetics we determine stability diagrams of spherical droplets as a function of dimensionless viscosity and reaction parameters. We determine concentration and flow fields inside and outside the droplets during shape changes and division. Our work shows that hydrodynamic flows tends to stabilize spherical shapes but that droplet division occurs for sufficiently strong chemical driving, sufficiently large droplet viscosity or sufficiently small surface tension. Active droplets could provide simple models for prebiotic protocells that are able to proliferate. Our work captures the key hydrodynamics of droplet division that could be observable in chemically active colloidal droplets

Statistics of Infima and Stopping Times of Entropy Production and Applications to Active Molecular Processes

We study the statistics of infima, stopping times and passage probabilities of entropy production in nonequilibrium steady states, and show that they are universal. We consider two examples of stopping times: first-passage times of entropy production and waiting times of stochastic processes, which are the times when a system reaches for the first time a given state. Our main results are: (i) the distribution of the global infimum of entropy production is exponential with mean equal to minus Boltzmann's constant; (ii) we find the exact expressions for the passage probabilities of entropy production to reach a given value; (iii) we derive a fluctuation theorem for stopping-time distributions of entropy production. These results have interesting implications for stochastic processes that can be discussed in simple colloidal systems and in active molecular processes. In particular, we show that the timing and statistics of discrete chemical transitions of molecular processes, such as, the steps of molecular motors, are governed by the statistics of entropy production. We also show that the extreme-value statistics of active molecular processes are governed by entropy production, for example, the infimum of entropy production of a motor can be related to the maximal excursion of a motor against the direction of an external force. Using this relation, we make predictions for the distribution of the maximum backtrack depth of RNA polymerases, which follows from our universal results for entropy-production infima.Comment: 30 pages, 13 figure

Sequential pattern formation governed by signaling gradients

Rhythmic and sequential segmentation of the embryonic body plan is a vital developmental patterning process in all vertebrate species. However, a theoretical framework capturing the emergence of dynamic patterns of gene expression from the interplay of cell oscillations with tissue elongation and shortening and with signaling gradients, is still missing. Here we show that a set of coupled genetic oscillators in an elongating tissue that is regulated by diffusing and advected signaling molecules can account for segmentation as a self-organized patterning process. This system can form a finite number of segments and the dynamics of segmentation and the total number of segments formed depend strongly on kinetic parameters describing tissue elongation and signaling molecules. The model accounts for existing experimental perturbations to signaling gradients, and makes testable predictions about novel perturbations. The variety of different patterns formed in our model can account for the variability of segmentation between different animal species.Comment: 12 pages, 5 figure

Casimir stresses in active nematic films

We calculate the Casimir stresses in a thin layer of active fluid with nematic order. By using a stochastic hydrodynamic approach for an active fluid layer of finite thickness L, we generalize the Casimir stress for nematic liquid crystals in thermal equilibrium to active systems. We show that the active Casimir stress differs significantly from its equilibrium counterpart. For contractile activity, the active Casimir stress, although attractive like its equilibrium counterpart, diverges logarithmically as L approaches a threshold of the spontaneous flow instability from below. In contrast, for small extensile activity, it is repulsive, has no divergence at any L and has a scaling with L different from its equilibrium counterpart

Droplet Ripening in Concentration Gradients

Living cells use phase separation and concentration gradients to organize chemical compartments in space. Here, we present a theoretical study of droplet dynamics in gradient systems. We derive the corresponding growth law of droplets and find that droplets exhibit a drift velocity and position dependent growth. As a consequence, the dissolution boundary moves through the system, thereby segregating droplets to one end. We show that for steep enough gradients, the ripening leads to a transient arrest of droplet growth that is induced by an narrowing of the droplet size distribution.Comment: 12 pages, 4 figure

Integral Fluctuation Relations for Entropy Production at Stopping Times

A stopping time $T$ is the first time when a trajectory of a stochastic process satisfies a specific criterion. In this paper, we use martingale theory to derive the integral fluctuation relation $\langle e^{-S_{\rm tot}(T)}\rangle=1$ for the stochastic entropy production $S_{\rm tot}$ in a stationary physical system at stochastic stopping times $T$. This fluctuation relation implies the law $\langle S_{\rm tot}(T)\rangle\geq 0$, which states that it is not possible to reduce entropy on average, even by stopping a stochastic process at a stopping time, and which we call the second law of thermodynamics at stopping times. This law implies bounds on the average amount of heat and work a system can extract from its environment when stopped at a random time. Furthermore, the integral fluctuation relation implies that certain fluctuations of entropy production are universal or are bounded by universal functions. These universal properties descend from the integral fluctuation relation by selecting appropriate stopping times: for example, when $T$ is a first-passage time for entropy production, then we obtain a bound on the statistics of negative records of entropy production. We illustrate these results on simple models of nonequilibrium systems described by Langevin equations and reveal two interesting phenomena. First, we demonstrate that isothermal mesoscopic systems can extract on average heat from their environment when stopped at a cleverly chosen moment and the second law at stopping times provides a bound on the average extracted heat. Second, we demonstrate that the average efficiency at stopping times of an autonomous stochastic heat engines, such as Feymann's ratchet, can be larger than the Carnot efficiency and the second law of thermodynamics at stopping times provides a bound on the average efficiency at stopping times.Comment: 37 pages, 6 figure

Generic Properties of Stochastic Entropy Production

We derive an Ito stochastic differential equation for entropy production in nonequilibrium Langevin processes. Introducing a random-time transformation, entropy production obeys a one-dimensional drift-diffusion equation, independent of the underlying physical model. This transformation allows us to identify generic properties of entropy production. It also leads to an exact uncertainty equality relating the Fano factor of entropy production and the Fano factor of the random time, which we also generalize to non steady-state conditions.Comment: 5 pages, 5 figures (contains Supplemental Material, 7 pages

Fluid pumping and active flexoelectricity can promote lumen nucleation in cell assemblies

We discuss the physical mechanisms that promote or suppress the nucleation of a fluid-filled lumen inside a cell assembly or a tissue. We discuss lumen formation in a continuum theory of tissue material properties in which the tissue is described as a two-fluid system to account for its permeation by the interstitial fluid, and we include fluid pumping as well as active electric effects. Considering a spherical geometry and a polarized tissue, our work shows that fluid pumping and tissue flexoelectricity play a crucial role in lumen formation. We furthermore explore the large variety of long-time states that are accessible for the cell aggregate and its lumen. Our work reveals a role of the coupling of mechanical, electrical and hydraulic phenomena in tissue lumen formation.Comment: Published versio

Cell body rocking is a dominant mechanism for flagellar synchronization in a swimming alga

The unicellular green algae Chlamydomonas swims with two flagella, which can synchronize their beat. Synchronized beating is required to swim both fast and straight. A long-standing hypothesis proposes that synchronization of flagella results from hydrodynamic coupling, but the details are not understood. Here, we present realistic hydrodynamic computations and high-speed tracking experiments of swimming cells that show how a perturbation from the synchronized state causes rotational motion of the cell body. This rotation feeds back on the flagellar dynamics via hydrodynamic friction forces and rapidly restores the synchronized state in our theory. We calculate that this `cell body rocking' provides the dominant contribution to synchronization in swimming cells, whereas direct hydrodynamic interactions between the flagella contribute negligibly. We experimentally confirmed the coupling between flagellar beating and cell body rocking predicted by our theory. This work appeared also in the Proceedings of the National Academy of Science of the U.S.A as: Geyer et al., PNAS 110(45), p. 18058(6), 2013.Comment: 40 pages, 15 color figure