arXiv subjects

Livio Nicola Carenza

Publications and source records attributed to Livio Nicola Carenza.

At least 19 recordsLinked to original sources

arXiv2025

Collective epithelial migration mediated by the unbinding of hexatic defects

Collective cell migration in epithelia relies on cell intercalation: a local remodelling of the cellular network that allows neighbouring cells to swap their positions. Unlike foams and passive cellular fluid, in epithelial intercalation these rearrangements crucially depend on activity. During these processes, the local geometry of the network and the contractile forces generated therein conspire to produce a burst of remodelling events, which collectively give rise to a vortical flow at the mesoscopic length scale. In this article we formulate a continuum theory of the mechanism driving this process, built upon recent advances towards understanding the hexatic (i.e. $6-$fold ordered) structure of epithelial layers. Using a combination of active hydrodynamics and cell-resolved numerical simulations, we demonstrate that cell intercalation takes place via the unbinding of topological defects, naturally initiated by fluctuations and whose late-times dynamics is governed by the interplay between passive attractive forces and active self-propulsion. Our approach sheds light on the structure of the cellular forces driving collective migration in epithelia and provides an explanation of the observed extensile activity of in vitro epithelial layers.

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arXiv2025

Activity drives self-assembly of passive soft inclusions in active nematics

Active nematics are out-of-equilibrium systems in which energy injection at the microscale drives emergent collective behaviors, from spontaneous flows to active turbulence. While the dynamics of these systems have been extensively studied, their potential for controlling the organization of embedded soft particles remains largely unexplored. Here, we investigate how passive droplets suspended in an active nematic fluid self-organize under varying activity levels and packing fractions. Through numerical simulations, we uncover a rich phase diagram featuring dynamic clustering, activity-induced gelation, and a novel inverse motility-induced phase separation regime where activity stabilizes dense droplet assemblies. Crucially, we demonstrate that temporal modulation of activity enables precise control over structural morphological transitions. Our results suggest new routes to design adaptive smart materials with tunable microstructure and dynamics, bridging active nematics with applications in programmable colloidal assembly and bio-inspired material design.

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arXiv2024

Thick liquid crystalline cholesteric shells

We numerically investigate the phase behavior of thick shells of cholesteric liquid crystals with tangential anchoring at the shell boundary. For achiral liquid crystal, we demonstrate a thickness-dependent transition from a configuration featuring four disclination line connecting the inner and outer surfaces to a state free of defect in the bulk, where each surface is topologically isolated and features two boojums. Incorporating chirality stabilizes novel defect arrangements, including a mixed state combining boojums and disclination lines and blue phases at high chirality and we demonstrate that shell thickness strongly modulates these transitions. Finally, we exploit the metastability features of the observed phases to obtain an elastically induced rearrangement of the shell surfaces during a cholesteric hysteresis cycle, stabilizing an alternative configuration that minimizes the free energy at low chirality. Our work paves the way for exploring dynamic behaviors under external fields, mixed anchoring conditions, or active flows.

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arXiv2024

Hydrodynamics and multiscale order in confluent epithelia

We formulate a hydrodynamic theory of confluent epithelia: i.e. monolayers of epithelial cells adhering to each other without gaps. Taking advantage of recent progresses toward establishing a general hydrodynamic theory of p-atic liquid crystals, we demonstrate that collectively migrating epithelia feature both nematic (i.e. p=2) and hexatic (i.e. p=6) order, with the former being dominant at large and the latter at small length scales. Such a remarkable multiscale liquid crystal order leaves a distinct signature in the system's structure factor, which exhibits two different power law scaling regimes, reflecting both the hexagonal geometry of small cells clusters, as well as the uniaxial structure of the global cellular flow. We support these analytical predictions with two different cell-resolved models of epithelia -- i.e. the self-propelled Voronoi model and the multiphase field model -- and highlight how momentum dissipation and noise influence the range of fluctuations at small length scales, thereby affecting the degree of cooperativity between cells. Our construction provides a theoretical framework to conceptualize the recent observation of multiscale order in layers of Madin-Darby canine kidney cells and pave the way for further theoretical developments.

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arXiv2023

Quasi long-ranged order in two-dimensional active liquid crystals

Quasi-long ranged order is the hallmark of two-dimensional liquid crystals. At equilibrium, this property implies that the correlation function of the local orientational order parameter decays with distance as a power law: i.e. $\sim|\boldsymbol{r}|^{-η_{p}}$, with $η_{p}$ a temperature-dependent exponent. While in general non-universal, $η_{p}=1/4$ universally at the Berezinskii-Kosterlitz-Thouless transition, where orientational order is lost because of the unbinding of disclinations. Motivated by recent experimental findings of liquid crystal order in confluent cell monolayers, here we demonstrate that, in {\em active} liquid crystals, the notion of quasi-long ranged order fundamentally differs from its equilibrium counterpart and is ultimately dictated by the interplay between translational and orientational dynamics. As a consequence, the exponent $η_{p}$ is allowed to vary in the range $0<η_{p}\le 2$, with the upper bound corresponding to the isotropic phase. Our theoretical predictions are supported by a survey of recent experimental data, reflecting a wide variety of different realization of orientational order in two dimensions.

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arXiv2023

Tuneable defect-curvature coupling and topological transitions in active shells

Recent experimental observations have suggested that topological defects can facilitate the creation of sharp features in developing embryos. Whereas these observations echo established knowledge about the interplay between geometry and topology in two-dimensional passive liquid crystals, the role of activity has mostly remained unexplored. In this article we focus on deformable shells consisting of either polar or nematic active liquid crystals and demonstrate that activity renders the mechanical coupling between defects and curvature much more involved and versatile than previously thought. Using a combination of linear stability analysis and three-dimensional computational fluid dynamics, we demonstrate that such a coupling can in fact be tuned, depending on the type of liquid crystal order, the specific structure of the defect (i.e. asters or vortices) and the nature of the active forces. In polar systems, this can drive a spectacular transition from spherical to toroidal topology, in the presence of large extensile activity. Our analysis strengthens the idea that defects could serve as topological morphogens and provides a number of predictions that could be tested in in vitro studies, for instance in the context of organoids.

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arXiv2023

Hydrodynamic Enhancement of $p$-atic Defect Dynamics

We investigate numerically and analytically the effects of hydrodynamics on the dynamics of topological defects in $p-$atic liquid crystals, i.e. two-dimensional liquid crystals with $p-$fold rotational symmetry. Importantly, we find that hydrodynamics fuels a generic passive self-propulsion mechanism for defects of winding number $s=(p-1)/p$ and arbitrary $p$. Strikingly, we discover that hydrodynamics always accelerates the annihilation dynamics of pairs of $\pm 1/p$ defects, and that, contrary to expectations, this effect increases with $p$. Our Letter paves the way towards understanding cell intercalation and other remodelling events in epithelial layers.

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arXiv2023

Topological Phases and Curvature-Driven Pattern Formation in Cholesteric Shells

We study the phase behaviour of cholesteric liquid crystal shells with different geometries. We compare the cases of tangential and no anchoring at the surface, focussing on the former case, which leads to a competition between the intrinsic tendency of the cholesteric to twist and the anchoring free energy which suppresses it. We then characterise the topological phases arising close to the isotropic-cholesteric transition. These typically consist of quasi-crystalline or amorphous tessellations of the surface by half-skyrmions, which are stable at lower and larger shell size respectively. For ellipsoidal shells, defects in the tessellation couple to local curvature, and according to the shell size they either migrate to the poles or distribute uniformly on the surface. For toroidal shells, the variations in the local curvature of the surface stabilises heterogeneous phases where cholesteric or isotropic patterns coexist with hexagonal lattices of half-skyrmions.

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arXiv2022

Epithelia are multiscale active liquid crystals

Biological processes such as embryogenesis, wound healing and cancer progression, crucially rely on the ability of epithelial cells to coordinate their mechanical activity over length scales order of magnitudes larger than the typical cellular size. While regulated by various signalling pathways, it has recently become evident that this behavior can additionally hinge on a minimal toolkit of physical mechanisms, of which liquid crystal order is the most prominent example. Yet, experimental and theoretical studies have given so far inconsistent results in this respect: whereas nematic order is often invoked in the interpretation of experimental data, computational models have instead suggested that hexatic order could serve as a linchpin for collective migration in confluent cell layers. In this article we resolve this dilemma. Using a combination of in vitro experiments on Madin-Darby canine kidney cells (MDCK), numerical simulations and analytical work, we demonstrate that both nematic and hexatic order are, in fact, present in epithelial layers, with the former being dominant at large length scales and the latter at small length scales. In MDCK GII cells on uncoated glass, these different types of liquid crystal order crossover at a length scale of the order of ten cell sizes. Our work sheds light on the emergent organization of living matter, provides a new framework for deciphering the structure of epithelia, and paves the way toward a comprehensive and predictive mesoscopic theory of tissues.

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arXiv2022

Theory of defect-mediated morphogenesis

Growing experimental evidence indicates that topological defects could serve as organizing centers in the morphogenesis of tissues. Here, we provide a quantitative explanation for this phenomenon, rooted in the buckling theory of deformable active polar liquid crystals. Using a combination of linear stability analysis and computational fluid dynamics, we demonstrate that active layers, such as confined cell monolayers, are unstable to the formation of protrusions in the presence of disclinations. The instability originates from an interplay between the focusing of the elastic forces, mediated by defects, and the renormalization of the system's surface tension by the active flow. The posttransitional regime is also characterized by several complex morphodynamical processes, such as oscillatory deformations, droplet nucleation, and active turbulence. Our findings offer an explanation of recent observations on tissue morphogenesis and shed light on the dynamics of active surfaces in general.

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arXiv2022

Yield-stress transition in suspensions of deformable droplets

Yield-stress materials, which require a sufficiently large forcing to flow, are currently ill-understood theoretically. To gain insight into their yielding transition, here we study numerically the rheology of a suspension of deformable droplets under pressure-driven flow. We show that the suspension displays yield-stress behaviour, with the droplets remaining motionless when the applied body-force is below a critical value. In the non-flowing phase, droplets jam to form an amorphous structure, whereas they order in the flowing phase. Yielding is linked to a percolation transition in the contacts of droplet-droplet overlaps, and requires suitable wetting boundary conditions and strict conservation of the droplet area to exist. Close to the yielding transition, we find strong oscillations in the droplet motion which closely resemble those found experimentally in confined colloidal glasses under flow. We show that even when droplets are static the underlying solvent moves by permeation, so that the viscosity of the composite system is never truly infinite, and, as we discuss, its precise value ceases to be a bulk material property of the system.

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arXiv2021

Cholesteric shells: two-dimensional blue fog and finite quasicrystals

We study the phase behaviour of a quasi-two dimensional cholesteric liquid crystal shell. We characterise the topological phases arising close to the isotropic-cholesteric transition, and show that they differ in a fundamental way from those observed on a flat geometry. For spherical shells, we discover two types of quasi-two dimensional topological phases: finite quasicrystals and amorphous structures, both made up by mixtures of polygonal tessellations of half-skyrmions. These structures generically emerge instead of regular double twist lattices because of geometric frustration, which disallows a regular hexagonal tiling of curved space. For toroidal shells, the variations in the local curvature of the surface stabilises heterogeneous phases where cholesteric patterns coexist with hexagonal lattices of half-skyrmions. Quasicrystals, amorphous and heterogeneous structures could be sought experimentally by self assembling cholesteric shells on the surface of emulsion droplets.

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arXiv2021

Rheology of active emulsions with negative effective viscosity

We numerically study by lattice Boltzmann simulations the rheological properties of an active emulsion made of a suspension of an active polar gel embedded in an isotropic passive background. We find that the hexatic equilibrium configuration of polar droplets is highly sensitive to both active injection and external forcing and may either lead to asymmetric unidirectional states which break top-bottom symmetry or symmetric ones. In this latter case, for large enough activity, the system develops a shear-thickening regime at low shear rates. Importantly, for larger external forcing a regime with stable negative effective viscosity is found. Moreover, at intermediate activity a region of multistability is encountered and we show that a maximum entropy production principle holds in selecting the most favorable state.

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arXiv2021

Cascade or not cascade? Energy transfer and elastic effects in active nematics

We numerically study the multi-scale properties of a 2d active gel to address the energy transfer mechanism. We find that activity is able to excite long-ranged distortions of the nematic pattern giving rise to spontaneous laminar flows and to a chaotic regime by further increasing the rate of active energy injection. By means of a scale-to-scale spectral analysis we find that the gel is basically driven by the local balancing between active injection and viscous dissipation, without any signal of non-linear hydrodynamical transfer and turbulent cascades. Furthermore, elasticity may qualitatively play an important role by transferring energy from small to larger scales through nemato-hydrodynamic interactions.

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arXiv2020

Rotation and propulsion in 3d active chiral droplets

Chirality is a recurrent theme in the study of biological systems, in which active processes are driven by the internal conversion of chemical energy into work. Bacterial flagella, acto-myosin filaments and microtubule bundles are active systems which are also intrinsically chiral. Despite some exploratory attempt to capture the relations between chirality and motility, no intrinsically chiral system has ever been analyzed so far. To address this gap in knowledge, here we study the effects of internal active forces and torques on a three-dimensional droplet of cholesteric liquid crystal (CLC) embedded in an isotropic liquid. We consider tangential anchoring of the liquid crystal director at the droplet surface. Contrary to what happens in nematics, where moderate extensile activity leads to droplet rotation, cholesteric active droplets exhibit a lot more complex and variegated behaviors. We find that extensile force dipole activity stabilises complex defect configurations whose orbiting dynamics couples to thermodynamic chirality to propel screw-like droplet motion. Instead, dipolar torque activity may either tighten or unwind the cholesteric helix, and if tuned can power rotations with an oscillatory angular velocity of zero mean.

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arXiv2019

Lattice Boltzmann Methods and Active Fluids

We review the state of the art of active fluids with particular attention to hydrodynamic continuous models and to the use of Lattice Boltzmann Methods (LBM) in this field. We present the thermodynamics of active fluids, in terms of liquid crystals modelling adapted to describe large-scale organization of active systems, as well as other effective phenomenological models. We discuss how LBM can be implemented to solve the hydrodynamics of active matter, starting from the case of a simple fluid, for which we explicitly recover the continuous equations by means of Chapman-Enskog expansion. Going beyond this simple case, we summarize how LBM can be used to treat complex and active fluids. We then review recent developments concerning some relevant topics in active matter that have been studied by means of LBM: spontaneous flow, self-propelled droplets, active emulsions, rheology, active turbulence, and active colloids.

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arXiv2019

Rheology of active polar emulsions: from linear to unidirectional and unviscid flow, and intermittent viscosity

The rheological behaviour of an emulsion made of an active polar component and an isotropic passive fluid is studied by lattice Boltzmann methods. Different flow regimes are found by varying the values of shear rate and extensile activity (occurring, e.g., in microtubule-motor suspensions). By increasing activity, a first transition occurs from linear flow regime to spontaneous persistent unidirectional macro-scale flow, followed by another transition either to (low shear) intermittent flow regime with coexistence of states with positive, negative, and vanishing apparent viscosity, or to (high shear) symmetric shear thinning regime. The different behaviours can be explained in terms of the dynamics of the polarization field close to the walls. A maximum entropy production principle selects the most likely states in the intermittent regime.

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arXiv2019

Dynamically asymmetric and bicontinuous morphologies in active emulsions

The morphology of a mixture made of a polar active gel immersed in an isotropic passive fluid is studied numerically. Lattice Boltzmann method is adopted to solve the Navier-Stokes equation and coupled to a finite-difference scheme used to integrate the dynamic equations of the concentration and of the polarization of the active component. By varying the relative amounts of the mixture phases, different structures can be observed. In the contractile case, at moderate values of activity, elongated structures are formed when the active component is less abundant, while a dynamic emulsion of passive droplets in an active matrix is obtained for symmetric composition. When the active component is extensile, aster-like rotating droplets and a phase-separated pattern appear for asymmetric and symmetric mixtures, respectively. The relevance of space dimensions in the overall morphology is shown by studying the system in three dimensions in the case of extensile asymmetric mixtures where interconnected tube-like structures span the whole system.

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