Search

Unlike coffee and cream that homogenize when stirred, growing micro-organisms (e.g., bacteria, baker's yeast) can actively kill each other and avoid mixing. How do such antagonistic interactions impact the growth and survival of competing strains, while being spatially advected by turbulent flows? By using numerical simulations of a continuum model, we study the dynamics of two antagonistic strains that are dispersed by incompressible turbulent flows in two spatial dimensions. A key parameter is the ratio of the fluid transport time to that of biological reproduction, which determines the winning strain that ultimately takes over the whole population from an initial heterogeneous state. By quantifying the probability and mean time for fixation along with the spatial structure of concentration fluctuations, we demonstrate how turbulence raises the threshold for biological nucleation and antagonism suppresses flow-induced mixing by depleting the population at interfaces. Our work highlights the unusual biological consequences of the interplay of turbulent fluid flows with antagonistic population dynamics, with potential implications for marine microbial ecology and origins of biological chirality., Comment: 7 pages, 4 figures

The assembly and co-evolution of supermassive black holes (SMBH) and their host galaxy stellar population is a key open questions in galaxy evolution. Stellar mass ($M_\star$) and star formation rate (SFR), are inferred by modeling the spectral energy distribution (SED). For galaxies triggering SMBH activity, the active galactic nucleus (AGN) contaminates the light at all wavelengths, hampering the inference of galaxy parameters. Incomplete AGN templates can lead to systematic overestimates of the stellar mass, biasing our understanding of AGN-galaxy co-evolution. This challenge has gained further impetus with the advent of sensitive wide-area surveys with millions of luminous AGN, including by eROSITA, Euclid and LSST. We aim to estimate the accuracy and bias of AGN host galaxy parameters and improve upon existing techniques. This work makes two contributions: 1) a new SED fitting code, GRAHSP, with a flexible, empirically motivated AGN model including a power law continuum emission lines, a FeII forest and a flexible infrared torus. We verify that our model reproduces published X-ray to infrared SEDs of AGN to better than 20\% accuracy. A fully Bayesian fit with nested sampling includes uncertainties in the model and the data, making the inference highly robust. 2) we created a benchmark photometric dataset where pure quasars are merged with non-AGN pure galaxies into a hybrid (Chimera) object but with known galaxy and AGN properties. Comparing the true and retrieved $M_\star$, SFR and AGN luminosities shows that previous codes systematically over-estimate $M_\star$ and SFR by 0.5 dex with a wide scatter of 0.7 dex, at AGN luminosities above 10^44 erg/s. In contrast, GRAHSP shows no bias on $M_\star$ and SFR. GRAHSP also estimates more realistic uncertainties. GRAHSP enables characterization of the environmental conditions conducive to black hole growth. (abridged), Comment: accepted in A&A

Topological defects play a central role in the physics of many materials, including magnets, superconductors and liquid crystals. In active fluids, defects become autonomous particles that spontaneously propel from internal active stresses and drive chaotic flows stirring the fluid. The intimate connection between defect textures and active flow suggests that properties of active materials can be engineered by controlling defects, but design principles for their spatiotemporal control remain elusive. Here we propose a symmetry-based additive strategy for using elementary activity patterns, as active topological tweezers, to create, move and braid such defects. By combining theory and simulations, we demonstrate how, at the collective level, spatial activity gradients act like electric fields which, when strong enough, induce an inverted topological polarization of defects, akin to a negative susceptibility dielectric. We harness this feature in a dynamic setting to collectively pattern and transport interacting active defects. Our work establishes an additive framework to sculpt flows and manipulate active defects in both space and time, paving the way to design programmable active and living materials for transport, memory and logic., Comment: 11 pages (including Methods), 5 figures. Changed title and format, final version

Boundary conditions dictate how fluids, including liquid crystals, flow when pumped through a channel. Can boundary conditions also be used to control internally driven active fluids that generate flows spontaneously? By using numerical simulations and stability analysis we explore how surface anchoring of active agents at the boundaries and substrate drag can be used to rectify coherent flow of an active polar fluid in a 2D channel. Upon increasing activity, a succession of dynamical states is obtained, from laminar flow to vortex arrays to eventual turbulence, that are controlled by the interplay between the hydrodynamic screening length and the extrapolation length quantifying the anchoring strength of the orientational order parameter. We highlight the key role of symmetry in both flow and order and show that coherent laminar flow with net throughput is only possible for weak anchoring and intermediate activity. Our work demonstrates the possibility of controlling the nature and properties of active flows in a channel simply by patterning the confining boundaries., Comment: 12 pages, 9 figures

The growth and evolution of microbial populations is often subjected to advection by fluid flows in spatially extended environments, with immediate consequences for questions of spatial population genetics in marine ecology, planktonic diversity and origin of life scenarios. Here, we review recent progress made in understanding this rich problem in the simplified setting of two competing genetic microbial strains subjected to fluid flows. As a pedagogical example we focus on antagonsim, i.e., two killer microorganism strains, each secreting toxins that impede the growth of their competitors (competitive exclusion), in the presence of stationary fluid flows. By solving two coupled reaction-diffusion equations that include advection by simple steady cellular flows composed of characteristic flow motifs in two dimensions (2d), we show how local flow shear and compressibility effects can interact with selective advantage to have a dramatic influence on genetic competition and fixation in spatially distributed populations. We analyze several 1d and 2d flow geometries including sources, sinks, vortices and saddles, and show how simple analytical models of the dynamics of the genetic interface can be used to shed light on the nucleation, coexistence and flow-driven instabilities of genetic drops. By exploiting an analogy with phase separation with nonconserved order parameters, we uncover how these genetic drops harness fluid flows for novel evolutionary strategies, even in the presence of number fluctuations, as confirmed by agent-based simulations as well., Comment: 29 pages, 22 figures

Understanding the complex patterns in space-time exhibited by active systems has been the subject of much interest in recent times. Complementing this forward problem is the inverse problem of controlling active matter. Here we use optimal control theory to pose the problem of transporting a slender drop of an active fluid and determine the dynamical profile of the active stresses to move it with minimal viscous dissipation. By parametrizing the position and size of the drop using a low-order description based on lubrication theory, we uncover a natural ''gather-move-spread'' strategy that leads to an optimal bound on the maximum achievable displacement of the drop relative to its size. In the continuum setting, the competition between passive surface tension, and active controls generates richer behaviour with futile oscillations and complex drop morphologies that trade internal dissipation against the transport cost to select optimal strategies. Our work combines active hydrodynamics and optimal control in a tractable and interpretable framework, and begins to pave the way for the spatiotemporal manipulation of active matter., Comment: 8 pages, 4 figures, SI available upon request

The common handsaw can be converted into a bowed musical instrument capable of producing exquisitely sustained notes when its blade is appropriately bent. Acoustic modes localized at an inflection point are known to underlie the saw's sonorous quality, yet the origin of localization has remained mysterious. Here we uncover a topological basis for the existence of localized modes, that relies on and is protected by spatial curvature. By combining experimental demonstrations, theory and computation, we show how spatial variations in blade curvature control the localization of these trapped states, allowing the saw to function as a geometrically tunable high quality oscillator. Our work establishes an unexpected connection between the dynamics of thin shells and topological insulators, and offers a robust principle to design high quality resonators across scales, from macroscopic instruments to nanoscale devices, simply through geometry., Comment: 17 pages, 3 figures, SI available upon request

The buckling of thin elastic sheets is a classic mechanical instability that occurs over a wide range of scales. In the extreme limit of atomically thin membranes like graphene, thermal fluctuations can dramatically modify such mechanical instabilities. We investigate here the delicate interplay of boundary conditions, nonlinear mechanics and thermal fluctuations in controlling buckling of confined thin sheets under isotropic compression. We identify two inequivalent mechanical ensembles based on the boundaries at constant strain (isometric) or at constant stress (isotensional) conditions.Remarkably, in the isometric ensemble, boundary conditions induce a novel long-ranged nonlinear interaction between the local tilt of the surface at distant points. This interaction combined with a spontaneously generated thermal tension leads to a renormalization group description of two distinct universality classes for thermalized buckling, realizing a mechanical variant of Fisher-renormalized critical exponents. We formulate a complete scaling theory of buckling as an unusual phase transition with a size dependent critical point, and discuss experimental ramifications for the mechanical manipulation of ultrathin nanomaterials., Comment: 31 pages, 8 figures (included summary of results and comparison to experiments)

Active matter encompasses different nonequilibrium systems in which individual constituents convert energy into non-conservative forces or motion at the microscale. This review provides an elementary introduction to the role of topology in active matter through experimentally relevant examples. Here, the focus lies on topological defects and topologically protected edge modes with an emphasis on the distinctive properties they acquire in active media. These paradigmatic examples represent two physically distinct classes of phenomena whose robustness can be traced to a common mathematical origin: the presence of topological invariants. These invariants are typically integer numbers that cannot be changed by continuous deformations of the relevant order parameters or physical parameters of the underlying medium. We first explain the mechanisms whereby topological defects self propel and proliferate in active nematics, leading to collective states which can be manipulated by geometry and patterning. Possible implications for active microfluidics and biological tissues are presented. We then illustrate how the propagation of waves in active fluids and solids is affected by the presence of topological invariants characterizing their dispersion relations. We discuss the relevance of these ideas for the design of robotic metamaterials and the properties of active granular and colloidal systems. Open theoretical and experimental challenges are presented as future research prospects., Comment: 23 pages, 3 figures, 4 boxes

Active matter consists of units that generate mechanical work by consuming energy. Examples include living systems, such as assemblies of bacteria and biological tissues, biopolymers driven by molecular motors, and suspensions of synthetic self-propelled particles. A central question in the field is to understand and control the self-organization of active assemblies in space and time. Most active systems exhibit either spatial order mediated by interactions that coordinate the spatial structure and the motion of active agents or the temporal synchronization of individual oscillatory dynamics. The simultaneous control of spatial and temporal organization is more challenging and generally requires complex interactions, such as reaction-diffusion hierarchies or genetically engineered cellular circuits. Here, we report a novel and simple means to simultaneously control the spatial and temporal self-organization of bacterial active matter. By confining an active bacterial suspension and manipulating a single macroscopic parameter, namely the viscoelasticity of the suspending fluid, we have found that the bacterial fluid first self-organizes in space into a millimeter-scale rotating vortex; then displays temporal organization as the giant vortex switches its global chirality periodically with tunable frequency, reminiscent of a torsional pendulum - a self-driven one. Combining experiments with an active matter model, we explain this striking behavior in terms of the interplay between active forcing and viscoelastic stress relaxation. Our findings advance the understanding of bacterial behavior in complex fluids, and demonstrate experimentally for the first time that rheological properties can be harnessed to control active matter flows. Coupled with actuation, our tunable self-oscillating bacterial vortex may be used as a "clock" for locomotion of soft robots and microfluidic pumping., Comment: 29 pages with 4 figure including Methods and Extended Data (with 10 figures); SI available upon request (accepted version)

In equilibrium, disorder conspires with topological defects to redefine the ordered states of matter in systems as diverse as crystals, superconductors and liquid crystals. Far from equilibrium, however, the consequences of quenched disorder on active condensed matter remain virtually uncharted. Here, we reveal a state of strongly disordered active matter with no counterparts in equilibrium: a dynamical vortex glass. Combining high-content microfluidic experiments and theory, we show how colloidal flocks collectively cruise through disorder without relaxing the topological singularities of their flows. The resulting state is highly dynamical but the flow patterns, shaped by a finite density of frozen vortices, are stationary and exponentially degenerated. Quenched isotropic disorder acts as a random gauge field turning active liquids into dynamical vortex glasses. We argue that this robust mechanism should shape the collective dynamics of a broad class of disordered active matter, from synthetic active nematics to collections of living cells exploring heterogeneous media., Comment: 9 pages, 3 figures (corrected a Grant number)

Epithelial tissues play a fundamental role in various morphogenetic events during development and early embryogenesis. Although epithelial monolayers are often modeled as two-dimensional (2D) elastic surfaces, they distinguish themselves from conventional thin elastic plates in three important ways - the presence of an apical-basal polarity, spatial variability of cellular thickness, and their nonequilibrium active nature. Here, we develop a minimal continuum model of a planar epithelial tissue as an active elastic material that incorporates all these features. We start from a full three-dimensional (3D)description of the tissue and derive an effective 2D model that captures, through the curvature of the apical surface, both the apical-basal asymmetry and the spatial geometry of the tissue. Crucially, variations of active stresses across the apical-basal axis lead to active torques that can drive curvature transitions. By identifying four distinct sources of activity, we find that bulk active stresses arising from actomyosin contractility and growth compete with boundary active tensions due to localized actomyosin cables and lamellipodial activity to generate the various states spanning the morphospace of a planar epithelium. Our treatment hence unifies 3D shape deformations through the coupled mechanics of apical curvature change and in-plane expansion/contraction of substrate-adhered tissues. Finally, we discuss the implications of our results for some biologically relevant processes such as tissue folding at the onset of lumen formation., Comment: 12 pages, 8 figures (including Appendix), published version including more calculations without changing results

Topological defects play a prominent role in the physics of two-dimensional materials. When driven out of equilibrium in active nematics, disclinations can acquire spontaneous self-propulsion and drive self-sustained flows upon proliferation. Here we construct a general hydrodynamic theory for a two-dimensional active nematic interrupted by a large number of such defects. Our equations describe the flows and spatio-temporal defect chaos characterizing active turbulence, even close to the defect unbinding transition. At high activity, nonequilibrium torques combined with many-body screening cause the active disclinations to spontaneously break rotational symmetry forming a collectively moving defect ordered polar liquid. By recognizing defects as the relevant quasiparticle excitations, we construct a comprehensive phase diagram for two-dimensional active nematics. Using our hydrodynamic approach, we additionally show that activity gradients can act like "electric fields", driving the sorting of topological charge. This demonstrates the versatility of our continuum model and its relevance for quantifying the use of spatially inhomogeneous activity for controlling active flows and for the fabrication of active devices with targeted transport capabilities., Comment: 18 pages, 7 figures, additional explanation provided with results unchanged

We develop a geometric approach to understand the mechanics of perforated thin elastic sheets, using the method of strain-dependent image elastic charges. This technique recognizes the buckling response of a hole under external load as a geometrically tuned mechanism of stress relief. We use a diagonally pulled square paper frame as a model system to quantitatively test and validate our approach. Specifically, we compare nonlinear force-extension curves and global displacement fields in theory and experiment. We find a strong softening of the force response accompanied by curvature localization at the inner corners of the buckled frame. Counterintuitively, though in complete agreement with our theory, for a range of intermediate hole sizes, wider frames are found to buckle more easily than narrower ones. Upon extending these ideas to many holes, we demonstrate that interacting elastic image charges can provide a useful kirigami design principle to selectively relax stresses in elastic materials., Comment: 6 pages, 4 figures; SI: 2 pages, 1 figure

We formulate the statistical dynamics of topological defects in the active nematic phase, formed in two dimensions by a collection of self-driven particles on a substrate. An important consequence of the non-equilibrium drive is the spontaneous motility of strength +1/2 disclinations. Starting from the hydrodynamic equations of active nematics, we derive an interacting particle description of defects that includes active torques. We show that activity, within perturbation theory, lowers the defect-unbinding transition temperature, determining a critical line in the temperature-activity plane that separates the quasi-long-range ordered (nematic) and disordered (isotropic) phases. Below a critical activity, defects remain bound as rotational noise decorrelates the directed dynamics of +1/2 defects, stabilizing the quasi-long-range ordered nematic state. This activity threshold vanishes at low temperature, leading to a re-entrant transition. At large enough activity, active forces always exceed thermal ones and the perturbative result fails, suggesting that in this regime activity will always disorder the system. Crucially, rotational diffusion being a two-dimensional phenomenon, defect unbinding cannot be described by a simplified one-dimensional model., Comment: 15 pages (including SI), 4 figures. Significant technical improvements without changing the results

Collections of self-propelled particles that move persistently by continuously consuming free energy are a paradigmatic example of active matter. In these systems, unlike Brownian "hot colloids", the breakdown of detailed balance yields a continuous production of entropy at steady state, even for an ideal active gas. We quantify the irreversibility for a non-interacting active particle in two dimensions by treating both conjugated and time-reversed dynamics. By starting with underdamped dynamics, we identify a hidden rate of entropy production required to maintain persistence and prevent the rapidly relaxing momenta from thermalizing, even in the limit of very large friction. Additionally, comparing two popular models of self-propulsion with identical dissipation on average, we find that the fluctuations and large deviations in work done are markedly different, providing thermodynamic insight into the varying extents to which macroscopically similar active matter systems may depart from equilibrium., Comment: 9 pages including SI, 3 figures (published version)

The dramatic effect kirigami, such as hole cutting, has on the elastic properties of thin sheets invites a study of the mechanics of thin elastic frames under an external load. Such frames can be thought of as modular elements needed to build any kirigami pattern. Here we develop the technique of elastic charges to address a variety of elastic problems involving thin sheets with perforations, focusing on frames with sharp corners. We find that holes generate elastic defects (partial disclinations) which act as sources of geometric incompatibility. Numerical and analytic studies are made of three different aspects of loaded frames - the deformed configuration itself, the effective mechanical properties in the form of force-extension curves and the buckling transition triggered by defects. This allows us to understand generic kirigami mechanics in terms of a set of force-dependent elastic charges with long-range interactions., Comment: 13 pages, 13 figures

We consider a collection of self-driven apolar particles on a substrate that organize into an active nematic phase at sufficiently high density or low noise. Using the dynamical renormalization group, we systematically study the 2d fluctuating ordered phase in a coarse-grained hydrodynamic description involving both the nematic director and the conserved density field. In the presence of noise, we show that the system always displays only quasi-long ranged orientational order beyond a crossover scale. A careful analysis of the nonlinearities permitted by symmetry reveals that activity is dangerously irrelevant over the linearized description, allowing giant number fluctuations to persist though now with strong finite-size effects and a non-universal scaling exponent. Nonlinear effects from the active currents lead to power law correlations in the density field thereby preventing macroscopic phase separation in the thermodynamic limit., Comment: 17 pages, 5 figures

Active systems on curved geometries are ubiquitous in the living world. In the presence of curvature orientationally ordered polar flocks are forced to be inhomogeneous, often requiring the presence of topological defects even in the steady state due to the constraints imposed by the topology of the underlying surface. In the presence of spontaneous flow the system additionally supports long-wavelength propagating sound modes which get gapped by the curvature of the underlying substrate. We analytically compute the steady state profile of an active polar flock on a two-sphere and a catenoid, and show that curvature and active flow together result in symmetry protected topological modes that get localized to special geodesics on the surface (the equator or the neck respectively). These modes are the analogue of edge states in electronic quantum Hall systems and provide unidirectional channels for information transport in the flock, robust against disorder and backscattering., Comment: 15 pages, 6 figures

In vitro reconstituted active systems, such as the ATP-driven microtubule bundle suspension developed by the Dogic group, provide a fertile testing ground for elucidating the phenomenology of active liquid crystalline states. Controlling such novel phases of matter crucially depends on our knowledge of their material and physical properties. In this letter, we show that the rheological properties of an active nematic film can be probed by varying its hydrodynamic coupling to a bounding oil layer. Using the motion of disclinations as intrinsic tracers of the flow field and a hydrodynamic model, we obtain an estimate for the as of now unknown shear viscosity of the nematic film. Knowing this now provides us with an additional handle for robust and precision tunable control of the emergent dynamics of active fluids., Comment: 5 pages, 4 figures, a SI file, and a SI video