Cell Division and Motility Enable Hexatic Order in Biological Tissues.

Biological tissues transform between solid- and liquidlike states in many fundamental physiological events. Recent experimental observations further suggest that in two-dimensional epithelial tissues these solid-liquid transformations can happen via intermediate states akin to the intermediate hexatic phases observed in equilibrium two-dimensional melting. The hexatic phase is characterized by quasi-long-range (power-law) orientational order but no translational order, thus endowing some structure to an otherwise structureless fluid.

Design rules for controlling active topological defects.

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.

Geometric control of tilt transition dynamics in single-clamped thermalized elastic sheets.

We study the finite-temperature dynamics of thin elastic sheets in a single-clamped cantilever configuration. This system is known to exhibit a tilt transition at which the preferred mean plane of the sheet shifts from horizontal to a plane above or below the horizontal. The resultant thermally roughened two-state (up/down) system possesses rich dynamics on multiple timescales.

Active cell divisions generate fourfold orientationally ordered phase in living tissue.

Morphogenesis, the process through which genes generate form, establishes tissue-scale order as a template for constructing the complex shapes of the body plan. The extensive growth required to build these ordered substrates is fuelled by cell proliferation, which, naively, should destroy order. Understanding how active morphogenetic mechanisms couple cellular and mechanical processes to generate order-rather than annihilate it-remains an outstanding question in animal development.

The role of non-affine deformations in the elastic behavior of the cellular vertex model.

The vertex model of epithelia describes the apical surface of a tissue as a tiling of polygonal cells, with a mechanical energy governed by deviations in cell shape from preferred, or target, area, , and perimeter, . The model exhibits a rigidity transition driven by geometric incompatibility as tuned by the target shape index, . For with (6) the perimeter of a regular hexagon of unit area, a cell can simultaneously attain both the preferred area and preferred perimeter.

Active nematic defects in compressible and incompressible flows.

We study the dynamics of active nematic films on a substrate driven by active flows with or without the incompressible constraint. Through simulations and theoretical analysis, we show that arch patterns are stable in the compressible case, while they become unstable under the incompressibility constraint. For compressible flows at high enough activity, stable arches organize themselves into a smecticlike pattern, which induce an associated global polar ordering of +1/2 nematic defects.

Anomalous elasticity of a cellular tissue vertex model.

Vertex models, such as those used to describe cellular tissue, have an energy controlled by deviations of each cell area and perimeter from target values. The constrained nonlinear relation between area and perimeter leads to new mechanical response. Here we provide a mean-field treatment of a highly simplified model: a uniform network of regular polygons with no topological rearrangements.

Flow around topological defects in active nematic films.

We study the active flow around isolated defects and the self-propulsion velocity of defects in an active nematic film with both viscous dissipation (with viscosity ) and frictional damping with a substrate. The interplay between these two dissipation mechanisms is controlled by the hydrodynamic dissipation length that screens the flows. For an isolated defect, in the absence of screening from other defects, the size of the shear vorticity around the defect is controlled by the system size .

Spontaneous Tilt of Single-Clamped Thermal Elastic Sheets.

Very thin elastic sheets, even at zero temperature, exhibit nonlinear elastic response by virtue of their dominant bending modes. Their behavior is even richer at finite temperature. Here, we use molecular dynamics to study the vibrations of a thermally fluctuating two-dimensional elastic sheet with one end clamped at its zero-temperature length.

Fluctuations can induce local nematic order and extensile stress in monolayers of motile cells.

Recent experiments in various cell types have shown that two-dimensional tissues often display local nematic order, with evidence of extensile stresses manifest in the dynamics of topological defects. Using a mesoscopic model where tissue flow is generated by fluctuating traction forces coupled to the nematic order parameter, we show that the resulting tissue dynamics can spontaneously produce local nematic order and an extensile internal stress. A key element of the model is the assumption that in the presence of local nematic alignment, cells preferentially crawl along the nematic axis, resulting in anisotropy of fluctuations.

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.

Erratum: Geometric Frustration and Solid-Solid Transitions in Model 2D Tissue [Phys. Rev. Lett. 120, 268105 (2018)].

This corrects the article DOI: 10.1103/PhysRevLett.120.

Non-uniform curvature and anisotropic deformation control wrinkling patterns on tori.

We investigate wrinkling patterns in a tri-layer torus consisting of an expanding thin outer layer, an intermediate soft layer and an inner core with a tunable shear modulus, inspired by pattern formation in developmental biology, such as follicle pattern formation during the development of chicken embryos. We show from large-scale finite element simulations that hexagonal wrinkling patterns form for stiff cores whereas stripe wrinkling patterns develop for soft cores. Hexagons and stripes co-exist to form hybrid patterns for cores with intermediate stiffness.

Nonlinear mechanics of thin frames.

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.

Kirigami Mechanics as Stress Relief by Elastic Charges.

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 an 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.

Topology and ground-state degeneracy of tetrahedral smectic vesicles.

Chemical design of block copolymers makes it possible to create polymer vesicles with tunable microscopic structure. Here we focus on a model of a vesicle made of smectic liquid-crystalline block copolymers at zero temperature. The vesicle assumes a faceted tetrahedral shape and the smectic layers arrange in a stack of parallel straight lines with topological defects localized at the vertices.

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 nonequilibrium 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.

Geometric Frustration and Solid-Solid Transitions in Model 2D Tissue.

We study the mechanical behavior of two-dimensional cellular tissues by formulating the continuum limit of discrete vertex models based on an energy that penalizes departures from a target area A_{0} and a target perimeter P_{0} for the component cells of the tissue. As the dimensionless target shape index s_{0}=(P_{0}/sqrt[A_{0}]) is varied, we find a transition from a soft elastic regime for a compatible target perimeter and area to a stiffer nonlinear elastic regime frustrated by geometric incompatibility. We show that the ground state in the soft regime has a family of degenerate solutions associated with zero modes for the target area and perimeter.

Thermal crumpling of perforated two-dimensional sheets.

Thermalized elastic membranes without distant self-avoidance are believed to undergo a crumpling transition when the microscopic bending stiffness is comparable to kT, the scale of thermal fluctuations. Most potential physical realizations of such membranes have a bending stiffness well in excess of experimentally achievable temperatures and are therefore unlikely ever to access the crumpling regime. We propose a mechanism to tune the onset of the crumpling transition by altering the geometry and topology of the sheet itself.

Defect driven shapes in nematic droplets: analogies with cell division.

Building on the striking similarity between the structure of the spindle during mitosis in living cells and nematic textures in confined liquid crystals, we use a continuum model of two-dimensional nematic liquid crystal droplets to examine the physical aspects of cell division. The model investigates the interplay between bulk elasticity of the microtubule assembly, described as a nematic liquid crystal, and surface elasticity of the cell cortex, modeled as a bounding flexible membrane, in controlling cell shape and division. The centrosomes at the spindle poles correspond to the cores of the topological defects required to accommodate nematic order in a closed geometry.

Effects of scars on icosahedral crystalline shell stability under external pressure.

We study how the stability of spherical crystalline shells under external pressure is influenced by the defect structure. In particular, we compare stability for shells with a minimal set of topologically required defects to shells with extended defect arrays (grain boundary "scars" with nonvanishing net disclination charge). We perform both Monte Carlo and conjugate gradient simulations to compare how shells with and without scars deform quasistatically under external hydrostatic pressure.

Defect dynamics in active nematics.

Topological defects are distinctive signatures of liquid crystals. They profoundly affect the viscoelastic behaviour of the fluid by constraining the orientational structure in a way that inevitably requires global changes not achievable with any set of local deformations. In active nematic liquid crystals, topological defects not only dictate the global structure of the director, but also act as local sources of motion, behaving as self-propelled particles.

Topology and dynamics of active nematic vesicles.

Engineering synthetic materials that mimic the remarkable complexity of living organisms is a fundamental challenge in science and technology. We studied the spatiotemporal patterns that emerge when an active nematic film of microtubules and molecular motors is encapsulated within a shape-changing lipid vesicle. Unlike in equilibrium systems, where defects are largely static structures, in active nematics defects move spontaneously and can be described as self-propelled particles.

Crystalline particle packings on constant mean curvature (Delaunay) surfaces.

We investigate the structure of crystalline particle arrays on constant mean curvature (CMC) surfaces of revolution. Such curved crystals have been realized physically by creating charge-stabilized colloidal arrays on liquid capillary bridges. CMC surfaces of revolution, classified by Delaunay in 1841, include the 2-sphere, the cylinder, the vanishing mean curvature catenoid (a minimal surface), and the richer and less investigated unduloid and nodoid.

Defect annihilation and proliferation in active nematics.

Liquid crystals inevitably possess topological defect excitations generated through boundary conditions, through applied fields, or in quenches to the ordered phase. In equilibrium, pairs of defects coarsen and annihilate as the uniform ground state is approached. Here we show that defects in active liquid crystals exhibit profoundly different behavior, depending on the degree of activity and its contractile or extensile character.