arXiv subjects

M. Cristina Marchetti

Publications and source records attributed to M. Cristina Marchetti.

At least 55 records · Page 3Linked to original sources

arXiv2020

Viscoelastic control of spatiotemporal order in bacterial active matter

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.

cond-mat.soft↗

arXiv2020

Emergence of dynamic vortex glasses in disordered polar active fluids

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.

cond-mat.soft↗

arXiv2020

Nonreciprocity as a generic route to traveling states

We examine a non-reciprocally coupled dynamical model of a mixture of two diffusing species. We demonstrate that nonreciprocity, which is encoded in the model via antagonistic cross diffusivities, provides a generic mechanism for the emergence of traveling patterns in purely diffusive systems with conservative dynamics. In the absence of non-reciprocity, the binary fluid mixture undergoes a phase transition from a homogeneous mixed state to a demixed state with spatially separated regions rich in one of the two components. Above a critical value of the parameter tuning non-reciprocity, the static demixed pattern acquires a finite velocity, resulting in a state that breaks both spatial and time translational symmetry, as well as the reflection parity of the static pattern. We elucidate the generic nature of the transition to traveling patterns using a minimal model that can be studied analytically. Our work has direct relevance to nonequilibrium assembly in mixtures of chemically interacting colloids that are known to exhibit non-reciprocal effective interactions, as well as to mixtures of active and passive agents where traveling states of the type predicted here have been observed in simulations. It also provides insight on transitions to traveling and oscillatory states seen in a broad range of nonreciprocal systems with non-conservative dynamics, from reaction-diffusion and prey-predators models to multispecies mixtures of microorganisms with antagonistic interactions.

cond-mat.soft↗

arXiv2020

Multi-defect Dynamics in Active Nematics

Recent experiments and numerical studies have drawn attention to the dynamics of active nematics. Two-dimensional active nematics flow spontaneously and exhibit spatiotemporal chaotic flows with proliferation of topological defects in the nematic texture. It has been proposed that the dynamics of active nematics can be understood in terms of the dynamics of interacting defects, propelled by active stress. Previous work has derived effective equations of motion for individual defects as quasi-particles moving in the mean field generated by other defects, but an effective theory governing multi-defect dynamics has remained out of reach. In this paper, we examine the dynamics of 2D active nematics in the limit of strong order and overdamped compressible flow. The activity-induced defect dynamics is formulated as a perturbation of the manifold of quasi-static nematic textures explicitly parameterized by defect positions. This makes it possible to derive a set of coupled ordinary differential equations governing defect (and therefore texture) dynamics. Interestingly, because of the non-orthogonality of textures associated with individual defects, their motion is coupled through a position dependent ``collective mobility" matrix. In addition to the familiar active self-propulsion of the $+1/2$ defect, we obtain new collective effects of activity that can be interpreted in terms of non-central and non-reciprocal interactions between defects.

cond-mat.soft↗

arXiv2020

How does the extracellular matrix affect the rigidity of an embedded spheroid?

Cellularized tissue and polymer networks can both transition from floppy to rigid as a function of their control parameters, and, yet, the two systems often mechanically interact, which may affect their respective rigidities. To study this interaction, we consider a vertex model with interfacial tension (a spheroid) embedded in a spring network in two dimensions. We identify two regimes with different global spheroid shapes, governed by the pressure resulting from competition between interfacial tension and tension in the network. In the first regime, the tissue remains compact, while in the second, a cavitation-like instability leads to the emergence of gaps at the tissue-network interface. Intriguingly, compression of the tissue promotes fluidization, while tension promotes cellular alignment and rigidification, with the mechanisms driving rigidification differing on either side of the instability.

q-bio.CB↗

arXiv2020

Shape and size changes of adherent elastic epithelia

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.

cond-mat.soft↗

arXiv2020

Solid-Liquid Transition of Deformable and Overlapping Active Particles

Experiments and theory have shown that cell monolayers and epithelial tissues exhibit solid-liquid and glass-liquid transitions. These transitions are biologically relevant to our understanding of embryonic development, wound healing, and cancer. Current models typically consider purely two-dimensional monolayers with no overlaps between neighboring cells. In reality, overlaps are important, and they may be precursors of cell extrusion -- a key biophysical process to maintain homeostasis in epithelial tissues. Here, we use a multi-phase field model to study the solid-liquid transition in a confluent monolayer of deformable cells which can overlap. When cells overlap rather than deform, we find that the melting transition changes from continuous to discontinuous, and that there is an intermittent regime close to the transition, where solid and liquid states alternate over time. By studying the dynamics of $5$- and $7$-fold disclinations in the hexagonal lattice formed by the cell centers, we observe that these correlate with spatial fluctuations in the cellular overlap, and that cell extrusion tends to initiate near $5$-fold disclinations.

cond-mat.soft↗

arXiv2020

Small-scale demixing in confluent biological tissues

Surface tension governed by differential adhesion can drive fluid particle mixtures to sort into separate regions, i.e., demix. Does the same phenomenon occur in confluent biological tissues? We begin to answer this question for epithelial monolayers with a combination of theory via a vertex model and experiments on keratinocyte monolayers. Vertex models are distinct from particle models in that the interactions between the cells are shape-based, as opposed to distance-dependent. We investigate whether a disparity in cell shape or size alone is sufficient to drive demixing in bidisperse vertex model fluid mixtures. Surprisingly, we observe that both types of bidisperse systems robustly mix on large lengthscales. On the other hand, shape disparity generates slight demixing over a few cell diameters, a phenomenon we term micro-demixing. This result can be understood by examining the differential energy barriers for neighbor exchanges (T1 transitions). Experiments with mixtures of wild-type and E-cadherin-deficient keratinocytes on a substrate are consistent with the predicted phenomenon of micro-demixing, which biology may exploit to create subtle patterning. The robustness of mixing at large scales, however, suggests that despite some differences in cell shape and size, progenitor cells can readily mix throughout a developing tissue until acquiring means of recognizing cells of different types.

cond-mat.soft↗

arXiv2019

Hydrodynamics of Active Defects: from order to chaos to defect ordering

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.

cond-mat.soft↗

arXiv2019

Phase transition to large scale coherent structures in 2d active matter turbulence

The collective motion of microswimmers in suspensions induce patterns of vortices on scales that are much larger than the characteristic size of a microswimmer, attaining a state called bacterial turbulence. Hydrodynamic turbulence acts on even larger scales and is dominated by inertial transport of energy. Using an established modification of the Navier-Stokes equation that accounts for the small scale forcing of hydrodynamic flow by microswimmers, we study the properties of a dense supensions of microswimmers in two dimensions, where the conservation of enstrophy can drive an inverse cascade through which energy is accumulated on the largest scales. We find that the dynamical and statistical properties of the flow show a sharp transition to the formation of vortices at the largest length scale. The results show that 2d bacterial and hydrodynamic turbulence are separated by a subcritical phase transition.

physics.flu-dyn↗

arXiv2019

Condensate formation and multiscale dynamics in two-dimensional active suspensions

The collective effects of microswimmers in active suspensions result in active turbulence, a spatiotemporally chaotic dynamics at mesoscale, which is characterized by the presence of vortices and jets at scales much larger than the characteristic size of the individual active constituents. To describe this dynamics, Navier-Stokes-based one-fluid models driven by small-scale forces have been proposed. Here, we provide a justification of such models for the case of dense suspensions in two dimensions (2d). We subsequently carry out an in-depth numerical study of the properties of one-fluid models as a function of the active driving in view of possible transition scenarios from active turbulence to large-scale pattern, referred to as condensate, formation induced by the classical inverse energy cascade in Newtonian 2d turbulence. Using a one-fluid model it was recently shown (Linkmann et al., Phys. Rev. Lett. (in press)) that two-dimensional active suspensions support two non-equilibrium steady states, one with a condensate and one without, which are separated by a subcritical transition. Here, we report further details on this transition such as hysteresis and discuss a low-dimensional model that describes the main features of the transition through nonlocal-in-scale coupling between the small-scale driving and the condensate.

physics.flu-dyn↗

arXiv2019

Glassy Dynamics in Models of Confluent Tissue with Mitosis and Apoptosis

Recent work on particle-based models of tissues has suggested that any finite rate of cell division and cell death is sufficient to fluidize an epithelial tissue. At the same time, experimental evidence has indicated the existence of glassy dynamics in some epithelial layers despite continued cell cycling. To address this discrepancy, we quantify the role of cell birth and death on glassy states in confluent tissues using simulations of an active vertex model that includes cell motility, cell division, and cell death. Our simulation data is consistent with a simple ansatz in which the rate of cell-life cycling and the rate of relaxation of the tissue in the absence of cell cycling contribute independently and additively to the overall rate of cell motion. Specifically, we find that a glass-like regime with caging behavior indicated by subdiffusive cell displacements can be achieved in systems with sufficiently low rates of cell cycling.

cond-mat.soft↗

arXiv2019

A self-driven phase transition drives Myxococcus xanthus fruiting body formation

Combining high-resolution single cell tracking experiments with numerical simulations, we show that starvation-induced fruiting body (FB) formation in Myxococcus xanthus is a phase separation driven by cells that tune their motility over time. The phase separation can be understood in terms of cell density and a dimensionless Peclet number that captures cell motility through speed and reversal frequency. Our work suggests that M. xanthus take advantage of a self-driven non-equilibrium phase transition that can be controlled at the single cell level.

physics.bio-ph↗

arXiv2019

Swimmer suspensions on substrates: anomalous stability and long-range order

We present a comprehensive theory of the dynamics and fluctuations of a two-dimensional suspension of polar active particles in an incompressible fluid confined to a substrate. We show that, depending on the sign of a single parameter, a state with polar orientational order is anomalously stable (or anomalously unstable), with a nonzero relaxation (or growth) rate for angular fluctuations at zero wavenumber. This screening of the broken-symmetry mode in the stable state does lead to conventional rather than giant number fluctuations as argued by Bricard et al., Nature ${\bf 503}$, 95 (2013), but their bend instability in a splay-stable flock does not exist and the polar phase has long-range order in two dimensions. Our theory also describes confined three-dimensional thin-film suspensions of active polar particles as well as dense compressible active polar rods, and predicts a flocking transition without a banding instability

cond-mat.soft↗

arXiv2018

Continuum models of collective cell migration

Collective cell migration plays a central role in tissue development, morphogenesis, wound repair and cancer progression. With the growing realization that physical forces mediate cell motility in development and physiology, a key biological question is how cells integrate molecular activities for force generation on multicellular scales. In this review we discuss recent advances in modeling collective cell migration using quantitative tools and approaches rooted in soft matter physics. We focus on theoretical models of cell aggregates as continuous active media, where the feedback between mechanical forces and regulatory biochemistry gives rise to rich collective dynamical behavior. This class of models provides a powerful predictive framework for the physiological dynamics that underlies many developmental processes, where cells need to collectively migrate like a viscous fluid to reach a target region, and then stiffen to support mechanical stresses and maintain tissue cohesion.

physics.bio-ph↗

arXiv2018

Defect unbinding in active nematics

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.

cond-mat.soft↗

arXiv2018

Hidden entropy production and work fluctuations in an ideal active gas

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.

cond-mat.soft↗

arXiv2018

Curvature-dependent tension and tangential flows at the interface of motility-induced phases

Purely repulsive active particles spontaneously undergo motility-induced phase separation (MIPS) into condensed and dilute phases. Remarkably, the mechanical tension measured along the interface between these phases is negative. In equilibrium this would imply an unstable interface that wants to expand, but these out-of-equilibrium systems display long-time stability and have intrinsically stiff boundaries. Here, we study this phenomenon in detail using active Brownian particle simulations and a novel frame of reference. By shifting from the global (or laboratory) frame to a local frame that follows the dynamics of the phase boundary, we observe correlations between the local curvature of the interface and the measured value of the tension. Importantly, our analysis reveals the presence of sustained local tangential motion of particles within a surface layer in both the gas and the dense regions. The combined tangential current in the gas and self-shearing of the surface of the dense phase suggest a stiffening interface that redirects particles along itself to heal local fluctuations. These currents restore the otherwise wildly fluctuating interface through an out-of-equilibrium Marangoni effect. We discuss the implications of our observations on phenomenological models of interfacial dynamics.

cond-mat.soft↗