Limits of economy and fidelity for programmable assembly of size-controlled triply periodic polyhedra.

We propose and investigate an extension of the Caspar-Klug symmetry principles for viral capsid assembly to the programmable assembly of size-controlled triply periodic polyhedra, discrete variants of the Primitive, Diamond, and Gyroid cubic minimal surfaces. Inspired by a recent class of programmable DNA origami colloids, we demonstrate that the economy of design in these crystalline assemblies-in terms of the growth of the number of distinct particle species required with the increased size-scale (e.g., periodicity)-is comparable to viral shells. We further test the role of geometric specificity in these assemblies via dynamical assembly simulations, which show that conditions for simultaneously efficient and high-fidelity assembly require an intermediate degree of flexibility of local angles and lengths in programmed assembly. Off-target misassembly occurs via incorporation of a variant of disclination defects, generalized to the case of hyperbolic crystals. The possibility of these topological defects is a direct consequence of the very same symmetry principles that underlie the economical design, exposing a basic tradeoff between design economy and fidelity of programmable, size controlled assembly.

Feedback linking cell envelope stiffness, curvature, and synthesis enables robust rod-shaped bacterial growth.

Bacterial growth is remarkably robust to environmental fluctuations, yet the mechanisms of growth-rate homeostasis are poorly understood. Here, we combine theory and experiment to infer mechanisms by which Escherichia coli adapts its growth rate in response to changes in osmolarity, a fundamental physicochemical property of the environment. The central tenet of our theoretical model is that cell-envelope expansion is only sensitive to local information, such as enzyme concentrations, cell-envelope curvature, and mechanical strain in the envelope. We constrained this model with quantitative measurements of the dynamics of E. coli elongation rate and cell width after hyperosmotic shock. Our analysis demonstrated that adaptive cell-envelope softening is a key process underlying growth-rate homeostasis. Furthermore, our model correctly predicted that softening does not occur above a critical hyperosmotic shock magnitude and precisely recapitulated the elongation-rate dynamics in response to shocks with magnitude larger than this threshold. Finally, we found that, to coordinately achieve growth-rate and cell-width homeostasis, cells employ direct feedback between cell-envelope curvature and envelope expansion. In sum, our analysis points to cellular mechanisms of bacterial growth-rate homeostasis and provides a practical theoretical framework for understanding this process.

Curvature screening in draped mechanical metamaterial sheets.

We develop a framework to understand the mechanics of metamaterial sheets on curved surfaces. Here we have constructed a continuum elastic theory of mechanical metamaterials by introducing an auxiliary, scalar gauge-like field that absorbs the strain along the soft mode and projects out the stiff ones. We propose a general form of the elastic energy of a mechanism based metamaterial sheet and specialize to the cases of dilational metamaterials and shear metamaterials conforming to positively and negatively curved substrates in the Föppl-Von Kármán limit of small strains. We perform numerical simulations of these systems and obtain good agreement with our analytical predictions. This work provides a framework that can be easily extended to explore non-linear soft modes in metamaterial elasticity in future.

Hidden symmetries generate rigid folding mechanisms in periodic origami.

We consider the zero-energy deformations of periodic origami sheets with generic crease patterns. Using a mapping from the linear folding motions of such sheets to force-bearing modes in conjunction with the Maxwell-Calladine index theorem we derive a relation between the number of linear folding motions and the number of rigid body modes that depends only on the average coordination number of the origami's vertices. This supports the recent result by Tachi [T. Tachi, Origami 6, 97-108 (2015)] which shows periodic origami sheets with triangular faces exhibit two-dimensional spaces of rigidly foldable cylindrical configurations. We also find, through analytical calculation and numerical simulation, branching of this configuration space from the flat state due to geometric compatibility constraints that prohibit finite Gaussian curvature. The same counting argument leads to pairing of spatially varying modes at opposite wavenumber in triangulated origami, preventing topological polarization but permitting a family of zero-energy deformations in the bulk that may be used to reconfigure the origami sheet.

Thermal Fluctuations of Singular Bar-Joint Mechanisms.

A bar-joint mechanism is a deformable assembly of freely rotating joints connected by stiff bars. Here we develop a formalism to study the equilibration of common bar-joint mechanisms with a thermal bath. When the constraints in a mechanism cease to be linearly independent, singularities can appear in its shape space, which is the part of its configuration space after discarding rigid motions. We show that the free-energy landscape of a mechanism at low temperatures is dominated by the neighborhoods of points that correspond to these singularities. We consider two example mechanisms with shape-space singularities and find that they are more likely to be found in configurations near the singularities than others. These findings are expected to help improve the design of nanomechanisms for various applications.

Robust folding of elastic origami.

Self-folding origami, structures that are engineered flat to fold into targeted, three-dimensional shapes, have many potential engineering applications. Though significant effort in recent years has been devoted to designing fold patterns that can achieve a variety of target shapes, recent work has also made clear that many origami structures exhibit multiple folding pathways, with a proliferation of geometric folding pathways as the origami structure becomes complex. The competition between these pathways can lead to structures that are programmed for one shape, yet fold incorrectly. To disentangle the features that lead to misfolding, we introduce a model of self-folding origami that accounts for the finite stretching rigidity of the origami faces and allows the computation of energy landscapes that lead to misfolding. We find that, in addition to the geometrical features of the origami, the finite elasticity of the nearly-flat origami configurations regulates the proliferation of potential misfolded states through a series of saddle-node bifurcations. We apply our model to one of the most common origami motifs, the symmetric "bird's foot," a single vertex with four folds. We show that though even a small error in programmed fold angles induces metastability in rigid origami, elasticity allows one to tune resilience to misfolding. In a more complex design, the "Randlett flapping bird," which has thousands of potential competing states, we further show that the number of actual observed minima is strongly determined by the structure's elasticity. In general, we show that elastic origami with both stiffer folds and less bendable faces self-folds better.

Mechanics of Metric Frustration in Contorted Filament Bundles: From Local Symmetry to Columnar Elasticity.

Bundles of filaments are subject to geometric frustration: certain deformations (e.g., bending while twisted) require longitudinal variations in spacing between filaments. While bundles are common-from protein fibers to yarns-the mechanical consequences of longitudinal frustration are unknown. We derive a geometrically nonlinear formalism for bundle mechanics, using a gaugelike symmetry under reptations along filament backbones. We relate force balance to orientational geometry and assess the elastic cost of frustration in twisted-toroidal bundles.

Topology in Nonlinear Mechanical Systems.

Many advancements have been made in the field of topological mechanics. The majority of the work, however, concerns the topological invariant in a linear theory. In this Letter, we present a generic prescription to define topological indices that accommodates nonlinear effects in mechanical systems without taking any approximation. Invoking the tools of differential geometry, a Z-valued quantity in terms of a topological index in differential geometry known as the Poincaré-Hopf index, which features the topological invariant of nonlinear zero modes (ZMs), is predicted. We further identify one type of topologically protected solitons that are robust to disorders. Our prescription constitutes a new direction of searching for novel topologically protected nonlinear ZMs in the future.

Theory and practice of origami in science.

No longer just the purview of artists and enthusiasts, origami engineering has emerged as a potentially powerful tool to create three dimensional structures on disparate scales. Whether origami (and the closely related kirigami) engineering can emerge as a useful technology will depend crucially on both fundamental theoretical advances as well as the development of further fabrication tools.

Distortion-controlled isotropic swelling: numerical study of free boundary swelling patterns.

Modern fabrication tools have now provided a number of platforms for designing flat sheets that, by virtue of their nonuniform growth, can buckle and fold into target three-dimensional structures. Theoretically, there is an infinitude of growth patterns that can produce the same shape, yet almost nothing is understood about which of these many growth patterns is optimal from the point of view of experiment, and few can even be realized at all. Here, we ask the question: what is the optimal way to design isotropic growth patterns for a given target shape? We propose a computational algorithm to produce optimal growth patterns by introducing cuts into the target surfaces. Within this framework, we propose that the patterns requiring the fewest or shortest cuts produce the best approximations to the target shape at finite thickness. The results are tested by simulation on spherical surfaces, and new challenges are highlighted for surfaces with both positive and negative Gaussian curvatures.

Overcurvature induced multistability of linked conical frusta: how a 'bendy straw' holds its shape.

We study the origins of multiple mechanically stable states exhibited by an elastic shell comprising multiple conical frusta, a geometry common to reconfigurable corrugated structures such as 'bendy straws'. This multistability is characterized by mechanical stability of axially extended and collapsed states, as well as a partially inverted 'bent' state that exhibits stability in any azimuthal direction. To understand the origin of this behavior, we study how geometry and internal stress affect the stability of linked conical frusta. We find that tuning geometrical parameters such as the frustum heights and cone angles can provide axial bistability, whereas stability in the bent state requires a sufficient amount of internal pre-stress, resulting from a mismatch between the natural and geometric curvatures of the shell. We provide insight into the latter effect through curvature analysis during deformation using X-ray computed tomography (CT), and with a simple mechanical model that captures the qualitative behavior of these highly reconfigurable systems.

Geometrically controlled snapping transitions in shells with curved creases.

Curvature and mechanics are intimately connected for thin materials, and this coupling between geometry and physical properties is readily seen in folded structures from intestinal villi and pollen grains to wrinkled membranes and programmable metamaterials. While the well-known rules and mechanisms behind folding a flat surface have been used to create deployable structures and shape transformable materials, folding of curved shells is still not fundamentally understood. Shells naturally deform by simultaneously bending and stretching, and while this coupling gives them great stability for engineering applications, it makes folding a surface of arbitrary curvature a nontrivial task. Here we discuss the geometry of folding a creased shell, and demonstrate theoretically the conditions under which it may fold smoothly. When these conditions are violated we show, using experiments and simulations, that shells undergo rapid snapping motion to fold from one stable configuration to another. Although material asymmetry is a proven mechanism for creating this bifurcation of stability, for the case of a creased shell, the inherent geometry itself serves as a barrier to folding. We discuss here how two fundamental geometric concepts, creases and curvature, combine to allow rapid transitions from one stable state to another. Independent of material system and length scale, the design rule that we introduce here explains how to generate snapping transitions in arbitrary surfaces, thus facilitating the creation of programmable multistable materials with fast actuation capabilities.

Contractility in an extensile system.

Essentially all biology is active and dynamic. Biological entities autonomously sense, compute, and respond using energy-coupled ratchets that can produce force and do work. The cytoskeleton, along with its associated proteins and motors, is a canonical example of biological active matter, which is responsible for cargo transport, cell motility, division, and morphology. Prior work on cytoskeletal active matter systems showed either extensile or contractile dynamics. Here, we demonstrate a cytoskeletal system that can control the direction of the network dynamics to be either extensile, contractile, or static depending on the concentration of filaments or weak, transient crosslinkers through systematic variation of the crosslinker or microtubule concentrations. Based on these new observations and our previously published results, we created a simple one-dimensional model of the interaction of filaments within a bundle. Despite its simplicity, our model recapitulates the observed activities of our experimental system, implying that the dynamics of our finite networks of bundles are driven by the local filament-filament interactions within the bundle. Finally, we show that contractile phases can result in autonomously motile networks that resemble cells. Our results reveal a fundamentally important aspect of cellular self-organization: weak, transient interacting species can tune their interaction strength directly by tuning the local concentration to act like a rheostat. In this case, when the weak, transient proteins crosslink microtubules, they can tune the dynamics of the network to change from extensile to contractile to static. Our experiments and model allow us to gain a deeper understanding of cytoskeletal dynamics and provide an new understanding of the importance of weak, transient interactions to soft and biological systems.

Optimal wrapping of liquid droplets with ultrathin sheets.

Elastic sheets offer a path to encapsulating a droplet of one fluid in another that is different from that of traditional molecular or particulate surfactants. In wrappings of fluids by sheets of moderate thickness with petals designed to curl into closed shapes, capillarity balances bending forces. Here, we show that, by using much thinner sheets, the constraints of this balance can be lifted to access a regime of high sheet bendability that brings three major advantages: ultrathin sheets automatically achieve optimally efficient shapes that maximize the enclosed volume of liquid for a fixed area of sheet; interfacial energies and mechanical properties of the sheet are irrelevant within this regime, thus allowing for further functionality; and complete coverage of the fluid can be achieved without special sheet designs. We propose and validate a general geometric model that captures the entire range of this new class of wrapped and partially wrapped shapes.

Origami structures with a critical transition to bistability arising from hidden degrees of freedom.

Origami is used beyond purely aesthetic pursuits to design responsive and customizable mechanical metamaterials. However, a generalized physical understanding of origami remains elusive, owing to the challenge of determining whether local kinematic constraints are globally compatible and to an incomplete understanding of how the folded sheet's material properties contribute to the overall mechanical response. Here, we show that the traditional square twist, whose crease pattern has zero degrees of freedom (DOF) and therefore should not be foldable, can nevertheless be folded by accessing bending deformations that are not explicit in the crease pattern. These hidden bending DOF are separated from the crease DOF by an energy gap that gives rise to a geometrically driven critical bifurcation between mono- and bistability. Noting its potential utility for fabricating mechanical switches, we use a temperature-responsive polymer-gel version of the square twist to demonstrate hysteretic folding dynamics at the sub-millimetre scale.

Grayscale gel lithography for programmed buckling of non-Euclidean hydrogel plates.

Shape programmable materials capable of morphing from a flat sheet into controlled three dimensional (3D) shapes offer promise in diverse areas including soft robotics, tunable optics, and bio-engineering. We describe a simple method of 'grayscale gel lithography' that relies on a digital micromirror array device (DMD) to control the dose of ultraviolet (UV) light, and therefore the extent of swelling of a photocrosslinkable poly(N-isopropyl acrylamide) (PNIPAm) copolymer film, with micrometer-scale spatial resolution. This approach allows for effectively smooth profiles of swelling to be prescribed, enabling the preparation of buckled 3D shapes with programmed Gaussian curvature.

Nonuniform growth and topological defects in the shaping of elastic sheets.

We demonstrate that shapes with zero Gaussian curvature, except at singularities, produced by the growth-induced buckling of a thin elastic sheet are the same as those produced by the Volterra construction of topological defects in which edges of an intrinsically flat surface are identified. With this connection, we study the problem of choosing an optimal pattern of growth for a prescribed developable surface, finding a fundamental trade-off between optimal design and the accuracy of the resulting shape which can be quantified by the length along which an edge should be identified.

Geometric localization of waves on thin elastic structures.

We consider the localization of elastic waves in thin elastic structures with spatially varying curvature profiles, using a curved rod and a singly curved shell as concrete examples. Previous studies on related problems have broadly focused on the localization of flexural waves on such structures. Here, using the semiclassical WKB approximation for multicomponent waves, we show that in addition to flexural waves, extensional and shear waves also form localized, bound states around points where the absolute curvature of the structure has a minimum. We also see excellent agreement between our numerical experiments and the semiclassical results, which hinges on the vanishing of two extra phases that arise in the semiclassical quantization rule. Our findings open up novel ways to fine-tune the acoustic and vibrational properties of thin elastic structures and raise the possibility of introducing new phenomena not easily captured by effective models of flexural waves alone.

How cells wrap around virus-like particles using extracellular filamentous protein structures.

Nanoparticles, such as viruses, can enter cells via endocytosis. During endocytosis, the cell surface wraps around the nanoparticle to effectively eat it. Prior focus has been on how nanoparticle size and shape impacts endocytosis. However, inspired by the noted presence of extracellular vimentin affecting viral and bacteria uptake, as well as the structure of coronaviruses, we construct a computational model in which both the cell-like construct and the virus-like construct contain filamentous protein structures protruding from their surfaces. We then study the impact of these additional degrees of freedom on viral wrapping. We find that cells with an optimal density of filamentous extracellular components (ECCs) are more likely to be infected as they uptake the virus faster and use relatively less cell surface area per individual virus. At the optimal density, the cell surface folds around the virus, and folds are faster and more efficient at wrapping the virus than crumple-like wrapping. We also find that cell surface bending rigidity helps generate folds, as bending rigidity enhances force transmission across the surface. However, changing other mechanical parameters, such as the stretching stiffness of filamentous ECCs or virus spikes, can drive crumple-like formation of the cell surface. We conclude with the implications of our study on the evolutionary pressures of virus-like particles, with a particular focus on the cellular microenvironment that may include filamentous ECCs.