Chemically active droplets provide simple models for cell-like systems that can grow and divide. Such active droplet systems are driven away from thermodynamic equilibrium and turn over chemically, which corresponds to a simple metabolism. We consider two scenarios of non-equilibrium driving. First, droplets are driven via the system boundaries by external reservoirs that supply nutrient and remove waste (boundary-driven). Second, droplets are driven by a chemical energy provided by a fuel in the bulk (bulk-driven). For both scenarios, we discuss the conservation of energy and matter as well as the balance of entropy. We use conserved and non-conserved fields to analyze the non-equilibrium steady states of active droplets. Using an effective droplet model, we explore droplet stability and instabilities leading to droplet division. Our work reveals that droplet division occurs quite generally in active droplet systems. Our results suggest that life-like processes such as metabolism and division can emerge in simple non-equilibrium systems that combine the physics of phase separation and chemical reactions.
Laboratoire de Physique de l’École Normale Supérieure, ENS, Université PSL, CNRS, Sorbonne Université, Université Paris Cité（巴黎高等师范学院物理实验室，法兰西学院，巴黎文理研究大学，法国国家科学研究中心，索邦大学，巴黎西岱大学）；Max Planck Institute for the Physics of Complex Systems（马克斯·普朗克复杂系统物理研究所）；Center for Systems Biology Dresden（德累斯顿系统生物学中心）；Cluster of Excellence Physics of Life, TU Dresden（德累斯顿工业大学生命物理学卓越集群）；Faculty of Mathematics, Natural Sciences, and Materials Engineering: Institute of Physics, University of Augsburg（奥格斯堡大学数学、自然科学与材料工程学院：物理研究所）
We numerically investigate the morphology and disclination line dynamics of active nematic droplets in three dimensions. Although our model only incorporates the simplest possible form of achiral active stress, active nematic droplets display an unprecedented range of complex morphologies. For extensile activity finger-like protrusions grow at points where disclination lines intersect the droplet surface. For contractile activity, however, the activity field drives cup-shaped droplet invagination, run-and-tumble motion or the formation of surface wrinkles. This diversity of behaviour is explai…
It has been proposed that during the early steps in the origin of life, small droplets could have formed via the segregation of molecules from complex mixtures by phase separation. These droplets could have provided chemical reaction centers. However, whether these droplets could divide and propagate is unclear. Here we examine the behavior of droplets in systems that are maintained away from thermodynamic equilibrium by an external supply of energy. In these systems, droplets grow by the addition of droplet material generated by chemical reactions. Surprisingly, we find that chemically driven…
From the mitotic spindle up to tissues and biofilms, many biological systems behave as active droplets, which often break symmetry and change shape spontaneously. Here, I show that active nematic droplets can experience a fingering instability. I consider an active fluid that acquires nematic order through anchoring at the droplet interface, and I predict its morphological stability in terms of three dimensionless parameters: the anchoring angle, the penetration length of nematic order compared to droplet size, and an active capillary number. Droplets with extensile (contractile) stresses and …
Active forces occurring within cells can drive crucial biological processes that involve spontaneous organization and shape change, such as cell division. Motivated by recent in vitro experiments of nematic droplets of cytoskeletal filaments and motors that self-organize and divide, we present a minimal hydrodynamic model that combines the nonequilibrium kinetics of motor-filament interactions with equilibrium nematic phase separation. The motors organize within droplets and structure filaments into polarized aster defects. At large motor activity, they can even deform or divide the droplet, o…
We present numerical simulations of active fluid droplets immersed in an external fluid in 2-dimensions { using} an Immersed Boundary method to simulate the fluid droplet interface as a Lagrangian mesh. We present results from two example systems, firstly an active isotropic fluid boundary consisting of particles that can bind and unbind from the interface and generate surface tension gradients through active contractility. Secondly, a droplet filled with an active polar fluid with { homeotropic} anchoring at the droplet interface. These two systems demonstrate spontaneous symmetry breaking an…
Large protein complexes are assembled from protein subunits to form a specific structure. In our theoretic work, we propose that assembly into the correct structure could be reliably achieved through an assembly line with a specific sequence of assembly steps. Using droplet interfaces to position compartment boundaries, we show that an assembly line can be self organized by active droplets. As a consequence, assembly steps can be arranged spatially so that a specific order of assembly is achieved and incorrect assembly is strongly suppressed.
We propose a generic model for thin films and shallow drops of a polar active liquid that have a free surface and are in contact with a solid substrate. The model couples evolution equations for the film height and the local polarization profile in the form of a gradient dynamics supplemented with active stresses and fluxes. A wetting energy for a partially wetting liquid is incorporated allowing for motion of the liquid-solid-gas contact line. This gives a consistent basis for the description of drops of dense bacterial suspensions or compact aggregates of living cells on solid substrates. As…
We investigate the swim pressure exerted by non-chiral and chiral active particles on convex or concave circular boundaries. Active particles are modeled as non-interacting and non-aligning self-propelled Brownian particles. The convex and concave circular boundaries are used as models representing a fixed inclusion immersed in an active bath and a cavity (or container) enclosing the active particles, respectively. We first present a detailed analysis of the role of convex versus concave boundary curvature and of the chirality of active particles on their spatial distribution, chirality-induce…
Active droplets emit a chemical solute at their surface that modifies their local interfacial tension. They exploit the nonlinear coupling of the convective transport of solute to the resulting Marangoni flows to self-propel. Such swimming droplets are by nature anti-chemotactic and are repelled by their own chemical wake or their neighbours. The rebound dynamics resulting from pairwise droplet interactions was recently analysed in detail for purely head-on collisions using a specific bispherical approach. Here, we extend this analysis and propose a reduced model of a generic collision to char…
It has been suggested recently that growth and division of a protocell could be modeled by a chemically active droplet with simple chemical reactions driven by an external fuel supply. This model is called the continuum model. Indeed it's numerical simulation reveals a shape instability which results in droplet division into two smaller droplets of equal size resembling cell division [1]. In this paper, we investigate the reduced version of the continuum model, which is called the effective model. This model is studied both in the linear and nonlinear regime. First, we perform a linear stabili…
Chemically-active droplets exhibit complex avoiding trajectories. While heterogeneity is inevitable in active matter experiments, it is mostly overlooked in their modelling. Exploiting its geometric simplicity, we fully-resolve the head-on collision of two swimming droplets of different radii and demonstrate that even a small contrast in size critically conditions their collision and subsequent dynamics. We identify three fundamentally-different regimes. The resulting high sensitivity of pairwise collisions is expected to profoundly affect their collective dynamics.
The swimming of cells, far from any boundary, can arise in the absence of long-range liquid-crystalline order within the cytoplasm, but simple models of this effect are lacking. Here we present a two-dimensional model of droplet self-propulsion involving two scalar fields, representing the cytoplasm and a contractile cortex. An active stress results from coupling between these fields; self-propulsion results when rotational symmetry is spontaneously broken. The swimming speed is predicted, and shown numerically, to vary linearly with the activity parameter and with the droplet area fraction. T…
We study numerically the behaviour of a mixture of a passive isotropic fluid and an active polar gel, in the presence of a surfactant favouring emulsification. Focussing on parameters for which the underlying free energy favours the lamellar phase in the passive limit, we show that the interplay between nonequilibrium and thermodynamic forces creates a range of multifarious exotic emulsions. When the active component is contractile (e.g., an actomyosin solution), moderate activity enhances the efficiency of lamellar ordering, whereas strong activity favours the creation of passive droplets wit…
Active fluid droplets surrounded by oil can spontaneously develop circulatory flows. However, the dynamics of the surrounding oil and their influence on the active fluid remain poorly understood. To investigate interactions between the active fluid and the passive oil across their interface, kinesin-driven microtubule-based active fluid droplets were immersed in oil and compressed into a cylinder-like shape. The droplet geometry supported intradroplet circulatory flows, but the circulation was suppressed when the thickness of the oil layer surrounding the droplet decreased. Experiments with tr…
Bacteria on surfaces exhibit collective behaviors, such as active turbulence and active stresses, which result from their motility, growth, and interactions with their local surroundings. However, interfacial deformations on soft surfaces and liquid interfaces caused by active growth, particularly over long time scales, are not well understood. Here, we describe experimental observations on the emergence of tubular structures arising from the growth of rod-shaped bacteria at the interface of oil droplets in water. Using microfluidics and timelapse microscopy, the dimensions and extension rates…
Evaporation-Induced Pattern Formation and Wetting in Active Microtubule-Kinesin Droplets
Harvard University（哈佛大学）；Max Planck Institute for Dynamics and Self-Organization（马克斯·普朗克动力学与自组织研究所）；University of Surrey（萨里大学）；Universität Konstanz（康斯坦茨大学）
We discuss the flow field and propulsion velocity of active droplets, which are driven by body forces residing on a rigid gel. The latter is modelled as a porous medium which gives rise to permeation forces. In the simplest model, the Brinkman equation, the porous medium is characterised by a single length scale $\ell$ --the square root of the permeability. We compute the flow fields inside and outside of the droplet as well as the energy dissipation as a function of $\ell$. We furthermore show that there are optimal gel fractions, giving rise to maximal linear and rotational velocities. In th…
How systems are endowed with migration capacity is a fascinating question with implications ranging from the design of novel active systems to the control of microbial populations. Bacteria, which can be found in a variety of environments, have developed among the richest set of locomotion mechanisms both at the microscopic and collective levels. Here, we uncover experimentally a new mode of collective bacterial motility in humid environment through the depinning of bacterial droplets. While capillary forces are notoriously enormous at the bacterial scale, even capable of pinning water droplet…
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 d…
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…
Experiments indicate that microdroplets undergoing micellar solubilization in the bulk of surfactant solution may excite Marangoni flows and self-propel spontaneously. Surprisingly, self-propulsion emerges even when the critical micelle concentration is exceeded and the Marangoni effect should be saturated. To explain this, we propose a novel model of a dissolving active droplet that is based on two fundamental assumptions: (a) products of the solubilization may inhibit surfactant adsorption; (b) solubilization prevents the formation of a monolayer of surfactant molecules at the droplet interf…
Active droplets swim as a result of the nonlinear advective coupling of the distribution of chemical species they consume or release with the Marangoni flows created by their non-uniform surface distribution. Most existing models focus on the self-propulsion of a single droplet in an unbounded fluid, which arises when diffusion is slow enough (i.e. beyond a critical Péclet number, $\mbox{Pe}_c$). Despite its experimental relevance, the coupled dynamics of multiple droplets and/or collision with a wall remains mostly unexplored. Using a novel approach based on a moving fitted bispherical grid, …
The bacterium Myxoccocus xanthus produces multicellular protective droplets called fruiting bodies when starved. These structures form initially through the active dewetting of cells into surface-bound droplets, where substantial flows of the material are needed as the fruiting bodies grow and become round. These dynamics are followed by a primitive developmental process in which the fluid-like droplets of motile cells mature into mechanically-resilient mounds of non-motile spores that can resist significant mechanical perturbation from the external environment. To date, the mechanical propert…
Migrating cell clusters as active droplets with asymmetric shape and contact angles
Max Planck Institute for the Physics of Complex Systems（马克斯·普朗克复杂系统物理研究所）；Center for Systems Biology Dresden（德累斯顿系统生物学中心）；Cluster of Excellence Physics of Life, Technische Universität Dresden（德累斯顿工业大学生命物理卓越集群）；Departament de Física de la Matèria Condensada, Universitat de Barcelona（巴塞罗那大学凝聚态物理系）；Universitat de Barcelona Institute of Complex Systems (UBICS)（巴塞罗那大学复杂系统研究所）；Institució Catalana de Recerca i Estudis Avançats (ICREA)（加泰罗尼亚研究高级学院）
Non-reciprocal interactions between condensates in chemically active mixtures
Max Planck Institute for Dynamics and Self-Organization（马克斯·普朗克动力学与自组织研究所）；SISSA — International School for Advanced Studies and INFN（国际高等研究院（SISSA）和意大利国家核物理研究所（INFN））；Rudolf Peierls Centre for Theoretical Physics, University of Oxford（牛津大学鲁道夫·派尔斯理论物理中心）
The breakup of a fluid jet into droplets has long fascinated natural scientists, with early research dating back to the 19th century. Infinitesimal perturbations to a jet grow because of surface tension, which eventually leads to breakup of the jet into droplets (Rayleigh-Plateau instability). Although this classical phenomenon has long been studied, it is not clear how it is modified when the fluid is replaced by an active fluid. In this study, we investigate instabilities of a jet of an active fluid. The active fluid is modelled by a suspension of microswimmers that propel themselves by gene…
