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.

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.

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.

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.

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.

Mechanical signaling cascades.

Mechanical computing has seen resurgent interest recently owing to the potential to embed sensing and computation into new classes of programmable metamaterials. To realize this, however, one must push signals from one part of a device to another and do so in a way that can be reset robustly. We investigate the propagation of signals in a bistable mechanical cascade uphill in energy. By identifying a penetration length for perturbations, we show that signals can propagate uphill for finite distances and map out parameters for this to occur. Experiments on soft elastomers corroborate our results.

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.

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.

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.

Energetic rigidity. II. Applications in examples of biological and underconstrained materials.

This is the second paper devoted to energetic rigidity, in which we apply our formalism to examples in two dimensions: Underconstrained random regular spring networks, vertex models, and jammed packings of soft particles. Spring networks and vertex models are both highly underconstrained, and first-order constraint counting does not predict their rigidity, but second-order rigidity does. In contrast, spherical jammed packings are overconstrained and thus first-order rigid, meaning that constraint counting is equivalent to energetic rigidity as long as prestresses in the system are sufficiently small. Aspherical jammed packings on the other hand have been shown to be jammed at hypostaticity, which we use to argue for a modified constraint counting for systems that are energetically rigid at quartic order.

Energetic rigidity. I. A unifying theory of mechanical stability.

Rigidity regulates the integrity and function of many physical and biological systems. This is the first of two papers on the origin of rigidity, wherein we propose that "energetic rigidity," in which all nontrivial deformations raise the energy of a structure, is a more useful notion of rigidity in practice than two more commonly used rigidity tests: Maxwell-Calladine constraint counting (first-order rigidity) and second-order rigidity. We find that constraint counting robustly predicts energetic rigidity only when the system has no states of self-stress. When the system has states of self-stress, we show that second-order rigidity can imply energetic rigidity in systems that are not considered rigid based on constraint counting, and is even more reliable than shear modulus. We also show that there may be systems for which neither first- nor second-order rigidity imply energetic rigidity. The formalism of energetic rigidity unifies our understanding of mechanical stability and also suggests new avenues for material design.

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.

Apolipoprotein Mimetic Peptide Inhibits Neutrophil-Driven Inflammatory Damage via Membrane Remodeling and Suppression of Cell Lysis.

Neutrophils are crucial for host defense but are notorious for causing sterile inflammatory damage. Activated neutrophils in inflamed tissue can liberate histone H4, which was recently shown to perpetuate inflammation by permeating membranes via the generation of negative Gaussian curvature (NGC), leading to lytic cell death. Here, we show that it is possible to build peptides or proteins that cancel NGC in membranes and thereby suppress pore formation, and demonstrate that they can inhibit H4 membrane remodeling and thereby reduce histone H4-driven lytic cell death and resultant inflammation. As a demonstration of principle, we use apolipoprotein A-I (apoA-I) mimetic peptide apoMP1. X-ray structural studies and theoretical calculations show that apoMP1 induces nanoscopic positive Gaussian curvature (PGC), which interacts with the NGC induced by the N-terminus of histone H4 (H4n) to inhibit membrane permeation. Interestingly, we show that induction of PGC can inhibit membrane-permeating activity in general and "turn off" diverse membrane-permeating molecules besides H4n. In vitro experiments show an apoMP1 dose-dependent rescue of H4 cytotoxicity. Using a mouse model, we show that tissue accumulation of neutrophils, release of neutrophil extracellular traps (NETs), and extracellular H4 all strongly correlate independently with local tissue cell death in multiple organs, but administration of apoMP1 inhibits histone H4-mediated cytotoxicity and strongly prevents organ tissue damage.

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.

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.

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.

Enabling Robust Self-Folding Origami by Pre-Biasing Vertex Buckling Direction.

Self-folding is a powerful approach to fabricate materials with complex 3D forms and advanced properties using planar patterning steps, but suffers from intrinsic limitations in robustness due to the highly bifurcated nature of configuration space around the flat state. Here, a simple mechanism is introduced to achieve robust self-folding of microscale origami by separating actuation into two discrete steps using different thermally responsive hydrogels. First, the vertices are pre-biased to move in the desired direction from the flat state by selectively swelling one of the two hydrogels at high temperature. Subsequently, the creases are folded toward their target angles by activating swelling of the second hydrogel upon cooling to room temperature. Since each vertex can be individually programmed to move upward or downward, it is possible to robustly select the desired branch even in multi-vertex structures with reasonably high complexity. This strategy provides key new principles for designing shaping-morphing materials that avoid undesired distractor states, expanding their potential applications in areas such as soft robotics, sensors, mechanical metamaterials, and deployable devices.

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.