In order to purposefully navigate their environments, animals rely on precise coordination between their sensory and motor systems. The integrated performance of circuits for sensorimotor control may be analyzed by quantifying an animal’s motile behavior in defined sensory environments. Here, we analyze the ability of the nematode C. elegans to crawl isothermally in spatial thermal gradients by quantifying the trajectories of individual worms responding to defined spatiotemporal thermal gradients. We show that sensorimotor control during isothermal tracking may be summarized as a strategy in which the worm changes the curvature of its propulsive undulations in response to temperature changes measured at its head. We show that a concise mathematical model for this strategy for sensorimotor control is consistent with the exquisite stability of the worm’s isothermal alignment in spatial thermal gradients as well as its more complex trajectories in spatiotemporal thermal gradients.

Recent experiments and simulations have demonstrated that particle-covered fluid/fluid interfaces can exist in stable nonspherical shapes as a result of the steric jamming of the interfacially trapped particles. The jamming confers the interface with solidlike properties. We provide an experimental and theoretical characterization of the mechanical properties of these armored objects, with attention given to the two-dimensional granular state of the interface. Small inhomogeneous stresses produce a plastic response, while homogeneous stresses produce a weak elastic response. Shear-driven particle-scale rearrangements explain the basic threshold needed to obtain the near-perfect plastic deformation that is observed. Furthermore, the inhomogeneous stress state of the interface is exhibited experimentally by using surfactants to destabilize the particles on the surface. Since the interfacially trapped particles retain their individual characteristics, armored interfaces can be recognized as a kind of composite material with distinct chemical,structural,

When a viscoelastic fluid blob is stretched out into a thin horizontal filament, it sags and falls gradually under its own weight, forming a catenary-like structure that evolves dynamically. If the ends are brought together rapidly after stretching, the falling filament tends to straighten by rising. These two effects are strongly influenced by the elasticity of the fluid and yield qualitatively different behaviours from the case of a purely viscous filament analysed previously (Teichman & Mahadevan, J. Fluid Mech. vol. 478, 2003, p. 71). Starting from the bulk equations for the motion of a viscoelastic fluid, we derive a simplified equation for the dynamics of a viscoelastic filament and analyse this equation in some simple settings to explain our observations.

Cytoskeletal microtubules have been proposed to infl uence cell shape and mechanics based on their ability to resist large-scale compressive forces exerted by the surrounding contractile cytoskeleton. Consistent with this, cytoplasmic microtubules are often highly curved and appear buckled because of compressive loads. However, the results of in vitro studies suggest that microtubules should buckle at much larger length scales, withstanding only exceedingly small compressive forces. This discrepancy calls into question the structural role of microtubules, and highlights our lack of quantitative knowledge of the magnitude of the forces they experience and can withstand in living cells. We show that intracellular microtubules do bear large-scale compressive loads from a variety of physiological forces, but their buckling wavelength is reduced signifi cantly because of mechanical coupling to the surrounding elastic cytoskeleton. We quantitatively explain this behavior, and show that this coupling dramatically increases the compressive forces that microtubules can sustain, suggesting they can make a more signifi cant structural contribution to the mechanical behavior of the cell than previously thought possible

An elastic rod model of a protein-bound DNA loop is adapted for application in multi-scale simulations of protein-DNA complexes. The classical Kirchhoff system of equations which describes the equilibrium structure of the elastic loop is modified to account for the intrinsic twist and curvature, anisotropic bending properties, and electrostatic charge of DNA. The effects of bending anisotropy and electrostatics are studied for the DNA loop clamped by the lac repressor protein. For two possible lengths of the loop, several topologically different conformations are predicted and extensively analyzed over the broad range of model parameters describing DNA bending and electrostatic properties. The scope and applications of the model in already accomplished and in future multi-scale studies of protein-DNA complexes are discussed.

We investigate the dynamics of fracture in drying films of colloidal silica. Water loss quenches the nanoparticle dispersionsto form a liquid-saturated elastic network of particlesthat relieves drying-induced strain by cracking. These cracks display intriguing intermittent motion originating from the deformation of arrested crack tips and aging of the elastic network. The dynamics of a single crack exhibits a universal evolution, described by a balance of the driving elastic power with the sum of interfacial power and the viscous dissipation rate of flowing interstitial fluid.

We investigate the dynamic fracture of a close-packed monolayer of particles, or particle raft, floating at a liquid-gas interface induced by the localized addition of surfactant. Unusually for a two-dimensional solid, our experiments show that the speed of crack propagation here is not affected by the elastic properties of the raft. Instead it is controlled by the rate at which surfactant is advected to the crack tip by means of the induced Marangoni flows. Further, the velocity of propagation is not constant in time and the length of the crack scales as t 3=4. More broadly, this surfactant-induced rupture of interfacial rafts suggests ways to manipulate them for applications.

We study the undulatory instability of a straight crack front generated by peeling a flexible elastic plate from a thin elastomeric adhesive film. We show that there is a threshold for the onset of the instability that is dependent on the ratio of two lengthscales that arise naturally in the problem: the thickness of the film and an elastic length defined by the stiffness of the plate and that of the film. A linear stability analysis predicts that the wavelength of the instability scales linearly with the film thickness. Our results are qualitatively and quantitatively consistent with recent experiments, and show how crack fronts may lose stability due to a competition between bulk and surface effects in the presence of multiple length scales.

We consider a microscopic model for the failure of soft adhesives in tension based on ideas of bond rupture under dynamic loading. By focusing on adhesive failure under loading at constant velocity, we demonstrate that bimodal curves of stress against strain may occur due to effects of finite polymer chain or bond length and characterize the loading conditions under which such bimodal behavior is observed. The results of this analysis are in qualitative agreement with experiments performed on unconfined adhesives in which failure does not occur by cavitation.

When a thin film of initially hydrated gelatin is allowed to dry from the surface, superficial changes in the structure of the material affect the local mechanical properties of the drying region. If the film is simultaneously subjected to large strain deformation (above 20%), a periodic pattern of wrinkles appears on the surface of the gelatin along the length of the sample in the direction of the applied force. These wrinkles are uniformly distributed on the surface of the gelatin with a wavelength that is much smaller than the sample thickness, which changes with sample composition, aging time, and deformation rate. We investigate these patterns via in situ environmentalscanning electron microscopy (ESEM) and provide a theory for their origin.

When a paintbrush is dipped into a pot of paint and pulled out, surface tension forces cause the individual hairs in the brush to coalesce even as the brush becomes impregnated with paint. We study a simple model of this elastocapillary interaction in the context of the surface-tension-driven vertical rise of a liquid between two long flexible hydrophilic sheets that are held a small distance apart at one end. We provide an analytic theory for the static shapes of the sheets as well as the liquid rise height which is different from that of the classical law of Jurin, and show that our experiments are quantitatively consistent with the theory.

Isotropic-nematic and nematic-nematic transitions from a homogeneous suspension of high aspect ratio magnetic rods are studied for both Maier-Saupe and Onsager excluded volume potentials. Asymptotic analysis in the vicinity of critical points yields insight into the stability and type of polarized nematic states emanating from nonpolarized equilibrium states. This, in conjunction with recently published global numerical results, yields a unified picture of the bifurcation diagram and provides a convenient base state to study effects of external orienting fields.

A cell’s contents are organized by a scaffolding of microtubules. These long, thin polymers continuously grow and shrink, and the structures of two forms of the constituent protein provide clues to how this occurs.

We study the behavior of a naturally straight highly extensible elastic filament subjected to large extensional and twisting strains. We find that two different phases can coexist for a range of parameter values: the plectoneme and the solenoid. A simple theory based on a neo-Hookean model for the material of the filament and accounting for the slender geometry suffices to explain these observations, and leads to a phase diagram that is consistent with observations. Extension and relaxation experiments on these phases show the presence of large hysteresis loops and sawtoothlike force-displacement curves which are different for the plectoneme and the solenoid.

We study the lubrication of fluid-immersed soft interfaces and show that elastic deformation couples tangential and normal forces and thus generates lift. We consider materials that deform easily, due to either geometry
e.g., a shell or constitutive properties
e.g., a gel or a rubber, so that the effects of pressure and temperature on the fluid properties may be neglected. Four different system geometries are considered: a rigid cylinder moving parallel to a soft layer coating a rigid substrate; a soft cylinder moving parallel to a rigid substrate; a cylindrical shell moving parallel to a rigid substrate; and finally a cylindrical conforming journal bearing coated with a thin soft layer. In addition, for the particular case of a soft layer coating a rigid substrate, we consider both elastic and poroelastic material responses. For all these cases, we find the same generic behavior: there is an optimal combination of geometric and material parameters that maximizes the dimensionless normal force as a function of the softness parameter
= hydrodynamic pressure/elastic stiffness = surface deflection/ gap thickness, which characterizes the fluid-induced deformation of the interface. The corresponding cases for a spherical slider are treated using scaling concepts.

Objects that float at the interface between a liquid and a gas interact because of interfacial deformation and the effect of gravity. We highlight the crucial role of buoyancy in this interaction, which, for small particles, prevails over the capillary suction that often is assumed to be the dominant effect. We emphasize this point using a simple classroom demonstration, and then derive the physical conditions leading to mutual attraction or repulsion. We also quantify the force of interaction in particular instances and present a simple dynamical model of this interaction. The results obtained from this model are validated by comparison to experimental results for the mutual attraction of two identical spherical particles. We consider some of the applications of the effect that can be found in nature and the laboratory

We present a unified framework for understanding the compaction of colloidal gels under their own weight. The dynamics of the collapse are determined by the value of the gravitational stress
g, as compared to the yield stress
Y of the network. For
g <
Y, gels collapse poroelastically, and their rate of compression decays exponentially in time. For
g >
Y, the network eventually yields, leading to rapid settling. In both cases, the rate of collapse is backflow limited, while its overall magnitude is determined by a balance between gravitational stress and network elastic stress.

Current models for protrusive motility in animal cells focus on cytoskeleton-based mechanisms, where localized protrusion is driven by local regulation of actin biochemistry1–3. In plants and fungi, protrusion is driven primarily by hydrostatic pressure4–6. For hydrostatic pressure to drive localized protrusion in animal cells7,8, it would have to be locally regulated, but current models treating cytoplasm as an incompressible viscoelastic continuum9 or viscous liquid10 require that hydrostatic pressure equilibrates essentially instantaneously over the whole cell. Here, we use cell blebs as reporters of local pressure in the cytoplasm. When we locally perfuse blebbing cells with cortex-relaxing drugs to dissipate pressure on one side, blebbing continues on the untreated side, implying non-equilibration of pressure on scales of approximately 10 mm and 10 s. We can account for localization of pressure by considering the cytoplasm as a contractile, elastic network infiltrated by cytosol. Motion of the fluid relative to the network generates spatially heterogeneous transients in the pressure field, and can be described in the framework of poroelasticity11,12.

The controlled folding and unfolding of maps, space structures, wings, leaves, petals, and other foldable laminae is potentially complicated by the independence of individual folds; as their number increases, there is a combinatorial explosion in the number of folded possibilities. The artificially constructed Miura-ori (1) pattern, with a periodic array of geometrically and elastically coupled mountain and valley folds (Fig. 1A), circumvents this complication by allowing the entire structure to be folded or unfolded simultaneously. Making such a pattern is not easy, so it may be surprising to find an elegant natural counterpart that is a few hundred millennia old. In Fig. 1B, we show the different stages of the opening of a hornbeam leaf that starts life in its bud as a Miura-ori folded pattern (2). Similar structures arise in insect wings (3) and elsewhere in nature (4), suggesting that these origami patterns are a result of convergent design. This raises a question of mechanism: How might this spatial organization of folds be brought about?

Stiff thin fi lms on soft substrates are both ancient and commonplace in nature; for instance, animal skin comprises a stiff epidermis attached to a soft dermis. Although more recent and rare, artifi cial skins are increasingly used in a broad range of applications, including fl exible electronics1 , tunable diff raction gratings2,3, force spectroscopy in cells4 , modern metrology methods5 , and other devices6–8. Here we show that model elastomeric artifi cial skins wrinkle in a hierarchical pattern consisting of self-similar buckles extending over fi ve orders of magnitude in length scale, ranging from a few nanometres to a few millimetres. We provide a mechanism for the formation of this hierarchical wrinkling pattern, and quantify our experimental fi ndings with both computations and a simple scaling theory. Th is allows us to harness the substrates for applications. In particular, we show how to use the multigeneration-wrinkled substrate for separating particles based on their size, while simultaneously forming linear chains of monodisperse particles.

The rapid closure of the Venus flytrap (Dionaea muscipula) leaf in about 100 ms is one of the fastest movements in the plant kingdom. This led Darwin to describe the plant as “one of the most wonderful in the world”1 . The trap closure is initiated by the mechanical stimulation of trigger hairs. Previous studies2–7 have focused on the biochemical response of the trigger hairs to stimuli and quantified the propagation of action potentials in the leaves. Here we complement these studies by considering the post-stimulation mechanical aspects of Venus flytrap closure. Using high-speed video imaging, non-invasive microscopy techniques and a simple theoretical model, we show that the fast closure of the trap results from a snap-buckling instability, the onset of which is controlled actively by the plant. Our study identifies an ingenious solution to scaling up movements in non-muscular engines and provides a general framework for understanding nastic motion in plants.

We give a hydrodynamical explanation for the chaotic behaviour of a dripping faucet using the results of the stability analysis of a static pendant drop and a proper orthogonal decomposition (POD) of the complete dynamics. We find that the only relevant modes are the two classical normal forms associated with a saddle– node–Andronov bifurcation and a Shilnikov homoclinic bifurcation. This allows us to construct a hierarchy of reduced-order models including maps and ordinary differential equations which are able to qualitatively explain prior experiments and numerical simulations of the governing partial differential equations and provide an explanation for the complexity in dripping. We also provide a new mechanical analogue for the dripping faucet and a simple rationale for the transition from dripping to jetting modes in the flow from a faucet.

We consider two of the simplest problems associated with the packing of a naturally flat thin elastic sheet. Both problems involve packing the sheet into a hollow cylinder; the first considers the partial contact of a cylindrically curved sheet with a cylindrical surface, while the second considers the partial contact of a conically curved sheet with the edge of a cylindrical surface. In each case, we solve the free-boundary problems to determine the shape, response and stability of the confined surfaces. In particular, we show that an exact description of both the cylindrical and conical structures is given by solutions of the Elastica equation, allowing us to present a unified description of a large class of elastic developable surfaces. This includes what is possibly the simplest example of strain localization, occurring at a point and forming one of the constituent elements of a crumpled elastic sheet.

We present a theoretical and experimental study of a method for the determination of the adhesion strength between a thin elastomeric film bonded to a rigid substrate and a flexible plate in a geometry common in the peel test. In particular, we characterize the work of adhesion in terms of the length of an equilibrium crack, generated by a spacer of known thickness wedged between the flexible plate and adhesive film, and the elastic and geometric properties of the film and the plate. We treat both the limit of perfect bonding and that of perfect slippage at the interface of the adhesive film and the flexible plate. A series of experiments allow us to verify the theory quantitatively and thus validate our method, which ought to be of value in many technological situations.

Minute concentrations of suspended particles can dramatically alter the behavior of a drying droplet. After a period of isotropic shrinkage, similar to droplets of a pure liquid, these droplets suddenly buckle like an elastic shell. While linear elasticity is able to describe the morphology of the buckled droplets, it fails to predict the onset of buckling. Instead, we find that buckling is coincident with a stress-induced fluid to solid transition in a shell of particles at a droplet’s surface, occurring when attractive capillary forces overcome stabilizing electrostatic forces between particles.

This report describes the spontaneous folding of flat elastomeric sheets, patterned with magnetic dipoles, into free-standing, 3D objects that are the topological equivalents of spherical shells. The path of the self-assembly is determined by a competition between mechanical and magnetic interactions. The potential of this strategy for the fabrication of 3D electronic devices is demonstrated by generating a simple electrical circuit surrounding a spherical cavity.

We give an explanation for the onset of fluid-flow-induced flutter in a flag. Our theory accounts for the various physical mechanisms at work: the finite length and the small but finite bending stiffness of the flag, the unsteadiness of the flow, the added mass effect, and vortex shedding from the trailing edge. Our analysis allows us to predict a critical speed for the onset of flapping as well as the frequency of flapping. We find that in a particular limit corresponding to a low-density fluid flowing over a soft high-density flag, the flapping instability is akin to a resonance between the mode of oscillation of a rigid pivoted airfoil in a flow and a hinged-free elastic plate vibrating in its lowest mode.

Photoresponsive solids such as nematic photoelastomers can undergo large deformations induced by light absorbed into rodlike molecules which bend and disrupt liquid crystal order. Significant variation of photoabsorption through the solid leads to nonuniform elastic deformations such as bending of beams and plates and pitting of layers. Such effects are also found in the presence of inhomogeneous thermal or swelling fields in solids or gels. We analyze the small deflection limit of these problems and show that beams made of these materials can have two elastically neutral planes, and that plates of these materials have a typical saddle shape. We also give a scaling analysis of the elasticity of photoinduced mounds and pits and speculate on their applications.

We consider some basic principles of fluid-induced lubrication at soft interfaces. In particular, we quantify how a soft substrate changes the geometry of and the forces between surfaces sliding past each other. By considering the model problem of a symmetric nonconforming contact moving tangentially to a thin elastic layer, we determine the normal force in the small and large deflection limit, and show that there is an optimal combination of material and geometric properties which maximizes the normal force. Our results can be generalized to a variety of other geometries which show the same qualitative behavior. Thus, they are relevant in the elastohydrodynamic lubrication of soft elastic and poroelastic gels and shells, and in the context of biolubrication in cartilaginous joints.

We consider the dynamics of an elastic sheet lubricated by the flow of a thin layer of fluid that separates it from a rigid wall. By considering long wavelength deformations of the sheet, we derive an evolution equation for its motion, accounting for the effects of elastic bending, viscous lubrication, and body forces. We then analyze various steady and unsteady problems for the sheet, such as peeling, healing, levitating, and bursting, using a combination of numerical simulation and dimensional analysis. On the macroscale, we corroborate our theory with a simple experiment, and, on the microscale, we analyze an oscillatory valve that can transform a continuous stream of fluid into a series of discrete pulses.

The linear and nonlinear viscoelastic response of networks of cross-linked and bundled cytoskeletal filaments demonstrates remarkable scaling with both frequency and applied prestress, which helps elucidate the origins of the viscoelasticity. The frequency dependence of the shear modulus reflects the underlying single-filament relaxation dynamics for 0:1–10 rad=sec. Moreover, the nonlinear strain stiffening of such networks exhibits a universal form as a function of prestress; this is quantitatively explained by the full force-extension relation of single semiflexible filaments.

Inspired by the observation that many naturally occurring adhesives arise as textured thin films, we consider the displacement-controlled peeling of a flexible plate from an incision-patterned thin adhesive elastic layer. We find that crack initiation from an incision on the film occurs at a load much higher than that required to propagate it on a smooth adhesive surface; multiple incisions thus cause the crack to propagate intermittently. Microscopically, this mode of crack initiation and propagation in geometrically confined thin adhesive films is related to the nucleation of cavitation bubbles behind the incision which must grow and coalesce before a viable crack propagates. Our theoretical analysis allows us to rationalize these experimental observations qualitatively and quantitatively and suggests a simple design criterion for increasing the interfacial fracture toughness of adhesive films

Networks of cross-linked and bundled actin filaments are ubiquitous in the cellular cytoskeleton, but their elasticity remains poorly understood. We show that these networks exhibit exceptional elastic behavior that reflects the mechanical properties of individual filaments. There are two distinct regimes of elasticity, one reflectingbendingof single filaments and a second reflectingstretchingof entropic fluctuations of filament length. The mechanical stiffness can vary by several decades with small changes in cross-link concentration, and can increase markedly upon application of external stress. We parameterize the full range of behavior in a state diagram and elucidate its origin with a robust model.

We consider the dynamics of capillary attraction between an articulated train of rigid rods floating at a liquid–gas interface and a nearby wall. We then explain some of the phenomena that are a result of the strong anisotropy and the extended nature of the system, such as the lining up next to the walling in a ‘zippering’ motion that is observed and compare our results qualitatively with those of experiments.

The acrosomal process of the sperm of the horseshoe crab (Limulus polyphemus) is a unique crystalline actin bundle, consisting of multiple actin filaments cross-linked by the actin-bundling protein, scruin. For successful fertilization, the acrosomal bundle must penetrate through a 30 mm thick jelly coat surrounding the egg and thus it must be sufficiently stiff. Here, we present two measurements of the bending stiffness of a single crystalline bundle of actin. Results from these measurements indicate that the actin:scruin composite bundle has an average elastic modulus of 2 GPa, which is similar to that of a single actin filament, and a bending stiffness that is more than two orders of magnitude larger than that of a bundle of uncross-linked actin filaments due to stiffening by the scruin matrix.

The classical Kirchhoff elastic-rod model applied to DNA is extended to account for sequence-dependent intrinsic twist and curvature, anisotropic bending rigidity, electrostatic force interactions, and overdamped Brownian motion in a solvent. The zero-temperature equilibrium rod model is then applied to study the structural basis of the function of the lac repressor protein in the lac operon of Escherichia coli. The structure of a DNA loop induced by the clamping of two distant DNA operator sites by lac repressor is investigated and the optimal geometries for the loop of length 76 bp are predicted. Further, the mimicked binding of catabolite gene activator protein (CAP) inside the loop provides solutions that might explain the experimentally observed synergy in DNA binding between the two proteins. Finally, a combined Monte Carlo and Brownian dynamics solver for a worm-like chain model is described and a preliminary analysis of DNA loop-formation kinetics is presented.

Inspired by the locomotion of terrestrial limbless animals, we study the motion of a lubricated rod of a hydrogel on a soft substrate. We show that it is possible to mimic observed biological gaits by vibrating the substrate and by using a variety of mechanisms to break longitudinal and lateral symmetry. Our simple theory and experiments provide a unified view of the creeping, undulating, and inchworming gaits observed in limbless locomotion on land, all of which originate as symmetry-breaking bifurcations of a simple base state associated with periodic longitudinal oscillations of a slender gel. These ideas are therefore also applicable to technological situations that involve moving small, soft solids on substrates.

We consider the gravity-induced draping of a 3D object with a naturally flat, isotropic elastic sheet. As the size of the sheet increases, we observe the appearance of new folded structures of increasing complexity that arise because of the competition between elasticity and gravity. We analyze some of the simpler 3D structures by determining their shape and analyzing their response and stability and show that these structures can easily switch between a number of metastable configurations. For more complex draperies, we derive scaling laws for the appearance and disappearance of new length scales. Our results are consistent with commonplace observations of drapes and complement large-scale computations of draping by providing benchmarks. They also yield a qualitative guide to fashion design and virtual reality animation.

The organization of individual actin filaments into higher-order structures is controlled by actin-binding proteins (ABPs). Although the biological significance of the ABPs is well documented, little is known about how bundling and cross-linking quantitatively affect the microstructure and mechanical properties of actin networks. Here we quantify the effect of the ABP scruin on actin networks by using imaging techniques, cosedimentation assays, multiparticle tracking, and bulk rheology. We show how the structure of the actin network is modified as the scruin concentration is varied, and we correlate these structural changes to variations in the resultant network elasticity.

We study the collective behaviour of a close-packed monolayer of non-Brownian particles at a fluid-liquid interface. Such a particle raft forms a two-dimensional elastic solid and can support anisotropic stresses and strains, e.g. it buckles in uniaxial compression and cracks in tension. We characterise this solid in terms of Young’s modulus and Poisson ratio derived from simple theoretical considerations and show the validity of these estimates by using an experimental buckling assay to deduce Young’s modulus.

Soft slender structures are ubiquitous in natural and artificial systems, in active and passive settings and across scales, from polymers and flagella, to snakes and space tethers. In this paper, we demonstrate the use of a simple and practical numerical implementation based on the Cosserat rod model to simulate the dynamics of filaments that can bend, twist, stretch and shear while interacting with complex environments via muscular activity, surface contact, friction and hydrodynamics. We validate our simulations by solving a number of forward problems involving the mechanics of passive filaments and comparing them with known analytical results, and extend them to study instabilities in stretched and twisted filaments that form solenoidal and plectonemic structures. We then study active filaments such as snakes and other slender organisms by solving inverse problems to identify optimal gaits for limbless locomotion on solid surfaces and in bulk liquids.

We present a multiresolution approach to modeling complexes between protein and DNA that contain looped or coiled DNA. The approach combines a coarse-grained model of the DNA loop, based on the classical theory of elasticity, with an atom level model of proteins and proteinDNA interfaces based on molecular dynamics. The coarse-grained DNA description is controlled through the atom level protein description and vice versa. The feasibility of the resulting multiscale modeling approach is demonstrated for a protein-DNA complex in which a protein called the E. coli lac repressor forces DNA into a 76 base pair loop. The required simulation involves 230,000 atoms, a number that would triple if both protein and DNA loops were described at the atomic level.

Thin adhesive pads used to attach objects to each other often fail catastrophically. Here we consider the nature of failure of such a pad under loading parallel to the adhesive substrate. To determine the modes of failure of the pad and to understand what limits its load bearing capacity, we conduct experiments with finite pads composed of a soft adhesive layer with a stiff backing and load them parallel to the surface of adhesion. We find that two different peeling mechanisms emerge as a function of the slenderness of the adhesive pad: an interfacial peeling mechanism that starts close to the pulling end for very long pads, and an unstable curling mechanism that starts at the opposite end for relatively short pads. A minimal theoretical framework allows us to explain our observations and reveals the adhesive bond stiffness as a dominant parameter in defining the peeling mode. A phase diagram that delineates the different regimes of peeling modes brings our experiments and theory together. Our results suggest that unstable peeling by curling may be more common than previously thought, and could perhaps occur naturally in such examples as the gecko foot.

During the 5 s of the acrosome reaction of Limulus polyphemus sperm, a 60-m-long bundle of scruin-decorated actin filaments straightens from a coiled conformation and extends from the cell. To identify the motive force for this movement, we examined the possible sources of chemical and mechanical energy and show that the coil releases 1013 J of stored mechanical D strain energy, whereas chemical energy derived from calcium binding is 1015 J. These measurements indicate that the coiled actin bundle extends by a spring-based mechanism, which is distinctly different from the better known polymerization or myosin-driven processes, and that calcium initiates but does not power the reaction.

Carbon nanotubes and biological filaments each spontaneously assemble into kinked helices, rings, and ‘‘tennis racket’’ shapes due to competition between elastic and interfacial effects. We show that the slender geometry is a more important determinant of the morphology than any molecular details. Our mesoscopic continuum theory is capable of quantifying observations of these structures and is suggestive of their occurrence in other filamentous assemblies as well

The wrinkling of thin elastic sheets occurs over a range of length scales, from the fine scale patterns in substrates on which cells crawl to the coarse wrinkles seen in clothes. Motivated by the wrinkling of a stretched elastic sheet, we deduce a general theory of wrinkling, valid far from the onset of the instability, using elementary geometry and the physics of bending and stretching. Our main result is a set of simple scaling laws; the wavelength of the wrinkles

K1=4, where K is the stiffness due to an ‘‘elastic substrate’’ effect with a multitude of origins, and the amplitude of the wrinkle A

. These could form the basis of a highly sensitive quantitative wrinkling assay for the mechanical characterization of thin solid membranes.