Motivated by a freely suspended graphene and polymerized membranes in soft and biological matter we present a detailed study of a tensionless elastic sheet in the presence of thermal fluctuations and quenched disorder. The manuscript is based on an extensive draft dating back to 1993, that was circulated privately. It presents the general theoretical framework and calculational details of numerous results, partial forms of which have been published in brief Letters (Le Doussal and Radzihovsky 1992). The experimental realization of atom-thin graphene sheets has driven a resurgence in this fascinating subject, making our dated predictions and their detailed derivations timely. To this end we analyze the statistical mechanics of a generalized D-dimensional elastic "membrane" embedded in d dimensions using a self-consistent screening approximation (SCSA), that has proved to be unprecedentedly accurate in this system, exact in three complementary limits: d --> infinity, D --> 4, and D=d. Focusing on the critical "flat" phase, for a homogeneous two-dimensional membrane embedded in three dimensions, we predict its universal length-scale dependent roughness, elastic moduli exponents, and a universal negative Poisson ratio of -1/3. We also extend these results to short- and long-range correlated random heterogeneity, predicting a variety of glassy wrinkled membrane states. Finally, we also predict and analyze a continuous crumpling transition in a "phantom" elastic sheet. We hope that this detailed presentation of the SCSA theory will be useful for further theoretical developments and corresponding experimental investigations on freely suspended graphene.
We evaluate the effective Hamiltonian governing, at the optically resolved scale, the elastic properties of micro-manipulated membranes. We identify floppy, entropic-tense and stretched-tense regimes, representing different behaviors of the effective area-elasticity of the membrane. The corresponding effective tension depends on the microscopic parameters (total area, bending rigidity) and on the optically visible area, which is controlled by the imposed external constraints. We successfully compare our predictions with recent data on micropipette experiments.
Recent theoretical advances in elasticity of membranes following Helfrich's famous spontaneous curvature model are summarized in this review. The governing equations describing equilibrium configurations of lipid vesicles, lipid membranes with free edges, and chiral lipid membranes are presented. Several analytic solutions to these equations and their corresponding configurations are demonstrated.
We amplify previous arguments why mean curvature should be used as measure of integration in calculating the effective bending rigidity of fluid membranes subjected to a weak background curvature. The stiffening of the membrane by its fluctuations, recently derived for spherical shapes, is recovered for cylindrical curvature. Employing curvilinear coordinates, we then discuss stiffening for arbitrary shapes, confirm that the elastic modulus of Gaussian curvature is not renormalized in the presence of fluctuations, and show for the first time that any spontaneous curvature also remains unchange…
Fluid membranes endowed with tangent-plane order (TPO) such as tilt- and hexatic order afford unique soft matter systems for investigating the interplay between elasticity, shape, topology, and thermal fluctuations. Using the spin-connection formulation of membrane energy we obtain equa- tions of equilibrium together with free boundary conditions for ground states of such membranes. We extend the spin-connection formulation to smectic liquid crystals with TPO and show that for chiral smectics-C* this generalization leads to experimentally verifiable consequences for dispirations having topolog…
One of the most widely used methods for determination of the bending elasticity modulus of model lipid membranes is the analysis of the shape fluctuations of nearly spherical lipid vesicles. The theoretical basis of this analysis is given by Milner and Safran. In their theory the stretching effects are not considered. In the present study we generalized their approach including the stretching effects deduced after an application of statistical mechanics of vesicles.
Using computational and theoretical approaches, we investigate the snap-through transition of buckled graphene membranes. Our main interest is related to the possibility of using the buckled membrane as a plate of capacitor with memory (memcapacitor). For this purpose, we performed molecular-dynamics (MD) simulations and elasticity theory calculations of the up-to-down and down-to-up snap-through transitions for membranes of several sizes. We have obtained expressions for the threshold switching forces for both up-to-down and down-to-up transitions. Moreover, the up-to-down threshold switching…
A phenomenological Landau elasticity for the shape, dilation, and lipid-tilt of bilayer membranes is developed. The shape mode couples with the sum of the monolayers' tilt, while the dilation mode couples with the difference of the monolayers' tilts. Interactions among membrane inclusions within regular arrays are discussed. Inclusions modifying the membrane thickness and/or inducing a tilt-difference due to their convex or concave shape yield a dilation-induced attraction and a tilt-difference-induced repulsion. The resulting interaction can stabilize 2D crystal phases, with the possible coex…
We study the shape, elasticity and fluctuations of the recently predicted (cond-mat/9510172) and subsequently observed (in numerical simulations) (cond-mat/9705059) tubule phase of anisotropic membranes, as well as the phase transitions into and out of it. This novel phase lies between the previously predicted flat and crumpled phases, both in temperature and in its physical properties: it is crumpled in one direction, and extended in the other. Its shape and elastic properties are characterized by a radius of gyration exponent $ν$ and an anisotropy exponent $z$. We derive scaling laws for the…
The purpose of this paper is to study the shapes and stabilities of bio-membranes within the framework of exterior differential forms. After a brief review of the current status in theoretical and experimental studies on the shapes of bio-membranes, a geometric scheme is proposed to discuss the shape equation of closed lipid bilayers, the shape equation and boundary conditions of open lipid bilayers and two-component membranes, the shape equation and in-plane strain equations of cell membranes with cross-linking structures, and the stabilities of closed lipid bilayers and cell membranes. The k…
Vesicles prepared in water from a series of diblock copolymers and termed "polymersomes" are physically characterized. With increasing molecular weight $\bar{M}_n$, the hydrophobic core thickness $d$ for the self-assembled bilayers of polyethyleneoxide - polybutadiene (PEO-PBD) increases up to 20 $nm$ - considerably greater than any previously studied lipid system. The mechanical responses of these membranes, specifically, the area elastic modulus $K_a$ and maximal areal strain $α_c$ are measured by micromanipulation. As expected for interface-dominated elasticity, $K_a$ ($\simeq$ 100 $pN/nm$)…
Cellular membranes are elastic lipid bilayers that contain a variety of proteins, including ion channels, receptors, and scaffolding proteins. These proteins are known to diffuse in the plane of the membrane and to influence the bending of the membrane. Experiments have shown that lipid flow in the plane of the membrane is closely coupled with the diffusion of proteins. Thus, there is a need for a comprehensive framework that accounts for the interplay between these processes. Here, we present a theory for the coupled in-plane viscous flow of lipids, diffusion of transmembrane proteins, and el…
Cell-cell adhesion is established by specific binding of receptor and ligand proteins. The adhesion bonds attract each other and often aggregate into large clusters that are central to many biological processes. One possible origin of attractive interactions between adhesion bonds is the elastic response of the membranes to their deformation by the bonds. Here, we analyze these elasticity-mediated interactions using a novel mean-field approach. Analysis of systems at different densities of bonds, $ϕ$, reveals that the phase diagram exhibits a nearly-universal behavior when the temperature $T$ …
Energies and equilibrium equations for thin elastic plates are discussed, with emphasis on several issues pertinent to recent approaches in soft condensed matter. Consequences of choice of basis, choice of invariant strain measures, and of approximating material energies as purely geometric in nature, are detailed. Ambiguities in the definitions of energies based on small-strain expansions, and in a typical informal process of dimensional reduction, are noted. A simple example serves to demonstrate that a commonly used bending energy has undesirable features, and it is suggested that a new the…
We preset a computational study of bending models for the curvature elasticity of lipid bilayer membranes that are relevant for simulations of vesicles and red blood cells. We compute bending energy and forces on triangulated meshes and evaluate and extend four well established schemes for their approximation: Kantor and Nelson 1987, Phys. Rev. A 36, 4020, Jülicher 1996, J. Phys. II France 6, 1797, Gompper and Kroll 1996, J. Phys. I France 6, 1305, and Meyer et. al. 2003 in Visualization and Mathematics III, Springer, p35, termed A, B, C, D. We present a comparative study of these four schemes…
We use coarse-grained Monte Carlo simulations to study the elastic properties of charged membranes in solutions of monovalent and pentavalent counterions. The simulation results of the two cases reveal trends opposite to each other. The bending rigidity and projected area increase with the membrane charge density for monovalent counterions, while they decrease for the pentavalent ions. These observations can be related to the counterion screening of the lipid charges. While the monovalent counterions only weakly screen the Coulomb interactions, which implies a repulsive Coulomb system, the mul…
We show that a smectic in a disordered medium (e.g., aerogel) exhibits anomalous elasticity, with the compression modulus B(k) vanishing and the bend modulus K(k) diverging as k --> 0. In addition, the effective disorder develops long ranged correlations. These divergences are much stronger than those driven by thermal fluctuations in pure smectics, and are controlled by a zero temperature glassy fixed point, which we study in an $ε=5-d$ expansion. We discuss the experimental implications of these theoretical predictions.
One problem of non-crystalline condensed matter (soft matter) is creating the right equilibrium between elasticity and viscosity, referred to as viscoelasticity. Manifestations of that can be found in everyday live, where the viscoelasticity in a tire needs to be balanced so it is still flexible and can dissipate shock-energy, yet hard enough for energy-saving operation. Similarly, the cartilage in joints needs to absorb shocks while operating at low- level friction with high elasticity. Two such examples with a biological applicability are stiff membranes, which allow for the sliding of joint…
The entropy of a polymer confined in a curved surface and the elastic free energy of a membrane consisting of polymers are obtained by scaling analysis. It is found that the elastic free energy of the membrane has the form of the in-plane strain energy plus Helfrich's curvature energy [Z. Naturforsch. C \textbf{28}, 693 (1973)]. The elastic constants in the free energy are obtained by discussing two simplified models: one is the polymer membrane without in-plane strains and asymmetry between its two sides, which is the counterpart of quantum mechanics in curved surface [Jensen and Koppe, Ann. …
Continuum elastic models that account for membrane thickness variations are especially useful in the description of nanoscale deformations due to the presence of membrane proteins with hydrophobic mismatch. We show that terms involving the gradient and the Laplacian of the area per lipid are significant and must be retained in the effective Hamiltonian of the membrane. We reanalyze recent numerical data, as well as experimental data on gramicidin channels, in light of our model. This analysis yields consistent results for the term stemming from the gradient of the area per molecule. The order …
We investigate interactions between an elastic membrane and a substrate characterized by quenched positional disorder in the height function. We show that the positional disorder transforms the standard secondary DLVO minimum into two separate states: the hovering state characterized by a planar membrane at a finite separation from the interface and a pinned state where the membrane follows closely the asperities of the substrate and is as a consequence quite corrugated. The transition between the two states is continuous and depends on the parameters of the underlying DLVO potential as well a…
The damping or attenuation coefficient of sound waves in solids due to impurities scales with the wavevector to the fourth power, also known as Rayleigh scattering. In amorphous solids, Rayleigh scattering may be enhanced by a logarithmic factor although computer simulations offer conflicting conclusions regarding this enhancement and its microscopic origin. We present a tensorial replica field-theoretic derivation based on heterogeneous or fluctuating elasticity (HE), which shows that long-range (power-law) spatial correlations of the elastic constants, is the origin of the logarithmic enhanc…
We study anomalous elasticity in the tubule phases of nematic and smectic elastomer membranes, which are flat in one direction and crumpled in another. These phases share the same macroscopic symmetry properties including spontaneously-broken in-plane isotropy and hence belong to the same universality class. Below an upper critical value $D_c =3$ of the membranes' intrinsic dimension D, thermal fluctuations renormalize the elasticity with respect to elastic displacements along the tubule axis so that elastic moduli for compression along the tubule axis and for bending the tubule axis become le…
A theory of vibrational excitations based on power-law spatial correlations in the elastic constants (or equivalently in the internal stress) is derived, in order to determine the vibrational density of states $D(ω)$ of disordered solids. The results provide the first prediction of a boson peak in amorphous materials where spatial correlations in the internal stresses (or elastic constants) are of power-law form, as is often the case in experimental systems, leading to logarithmic enhancement of (Rayleigh) phonon attenuation. A logarithmic correction of the form $\sim -ω^{2}\lnω$ is predicted …
We have studied the electrostatic screening effect of NaCl solutions on the interactions between anionic lipid bilayers in the fluid lamellar phase using a Poisson-Boltzmann based mean-field approach with constant charge and constant potential limiting charge regulation boundary conditions. The full DLVO potential, including the electrostatic, hydration and van der Waals interactions, was coupled to thermal bending fluctuations of the membranes via a variational Gaussian Ansatz. This allowed us to analyze the coupling between the osmotic pressure and the fluctuation amplitudes and compare them…
We study the force generation by a set of parallel actin filaments growing against an elastic membrane. The elastic membrane tries to stay flat and any deformation from this flat state, either caused by thermal fluctuations or due to protrusive polymerization force exerted by the filaments, costs energy. We study two lattice models to describe the membrane dynamics. In one case, the energy cost is assumed to be proportional to the absolute magnitude of the height gradient (gradient model) and in the other case it is proportional to the square of the height gradient (Gaussian model). For the gr…
The properties of crystals consisting of several components can be widely tuned. Often solid solutions are produced, where substitutional or interstitional disorder determines the crystal thermodynamic and mechanical properties. The chemical and structural disorder impedes the study of the elasticity of such solid solutions, since standard procedures like potential expansions cannot be applied. We present a generalization of a density-functional based approach recently developed for one-component crystals to multi-component crystals. It yields expressions for the elastic constants valid in sol…
We have studied two types of meshwork models by using the canonical Monte Carlo simulation technique. The first meshwork model has elastic junctions, which are composed of vertices, bonds, and triangles, while the second model has rigid junctions, which are hexagonal (or pentagonal) rigid plates. Two-dimensional elasticity is assumed only at the elastic junctions in the first model, and no two-dimensional bending elasticity is assumed in the second model. Both of the meshworks are of spherical topology. We find that both models undergo a first-order collapsing transition between the smooth sph…
The adhesion dynamics of a membrane confined between two permeable walls is studied using a two-dimensional hydrodynamic model. The membrane morphology decomposes into adhesion patches on the upper and the lower walls and obeys a nonlinear evolution equation that resembles that of phase separation dynamics, which is known to lead to coarsening, i.e. to the endless growth of the adhesion patches. However, due to the membrane bending rigidity the system evolves towards a frozen state without coarsening. This frozen state exhibits an order-disorder transition when increasing the permeability of t…
Prestress in amorphous solids bears the memory of their formation, and plays a profound role in their mechanical properties, from stiffening or softening elastic moduli to shifting frequencies of vibrational modes, as well as directing yielding and solidification in the nonlinear regime. Here we develop a set of mathematical tools to investigate elasticity of prestressed discrete networks, which disentangles the effects from disorder in configuration and disorder in prestress. Applying these methods to prestressed triangular lattices and a computational model of amorphous solids, we demonstrat…
Chiral rodlike colloids exposed to strong depletion attraction may self-assemble into chiral membranes whose twisted director field differs from that of a 3D bulk chiral nematic. We formulate a simple microscopic variational theory to determine the elastic moduli of rods assembled into a bi-dimensional smectic membrane. The approach is based on a simple Onsager-Straley theory for a non-uniform director field that we apply to describe rod twist within the membrane. A microscopic approach enables a detailed estimate of the individual Frank elastic moduli (splay, twist and bend) as well as the tw…
The relation between elasticity and yielding is investigated in a model polymer solid by Molecular-Dynamics simulations. By changing the bending stiffness of the chain and the bond length, semicrystalline and disordered glassy polymers - both with bond disorder - as well as nematic glassy polymers with bond ordering are obtained. It is found that in systems with bond disorder the ratio Ty/G between the shear yield strength Ty and the shear modulus G is close to the universal value of the atomic metallic glasses. The increase of the local nematic order in glasses leads to the increase of the sh…
Adhesion between a T cell and an antigen presenting cell is achieved by TCR-pMHC and LFA1-ICAM1 protein complexes. These segregate to form a special pattern, known as the immunological synapse (IS), consisting of a central quasi-circular domain of TCR-pMHC bonds surrounded by a peripheral domain of LFA1-ICAM1 complexes. Insights gained from imaging studies had led to the conclusion that the formation of the central adhesion domain in the IS is driven by active (ATP-driven) mechanisms. Recent studies, however, suggested that passive (thermodynamic) mechanisms may also play an important role in …
We present a simple, and physically motivated, coarse-grained model of a lipid bilayer, suited for micron scale computer simulations. Each ~25 nm^2 patch of bilayer is represented by a spherical particle. Mimicking forces of hydrophobic association, multi-particle interactions suppress the exposure of each sphere's equator to its implicit solvent surroundings. The requirement of high equatorial density stabilizes two-dimensional structures without necessitating crystalline order, allowing us to match both the elasticity and fluidity of natural lipid membranes. We illustrate the model's versati…
Lipid monolayers and bilayers have been used as experimental models for the investigation of membrane thermal transitions. The main transition takes place near ambient temperatures for several lipids and reflects the order-disorder transition of lipid hydrocarbonic chains, which is accompanied by a small density gap. Equivalence between the transitions in the two systems has been argued by several authors. The two-state statistical model adopted by numerous authors for different properties of the membrane, such as permeability, diffusion, mixture or insertion of cholesterol or protein, is inad…
Open cellular solids usually possess random microstructures that may contain a characteristic length scale, such as the cell size. This gives rise to size dependent mechanical properties where large systems behave differently from small systems. Furthermore, these structures are often irregular, which not only affects the size dependent behaviour but also leads to significant property variations among different microstructure realizations. The computational model for cellular microstructures is based on networks of Timoshenko beams. It is a computationally efficient approach allowing to obtain…
We examine how the distribution of contour lengths and the high-stretch stiffening of individual chain segments affect the macroscopic shear modulus of flexible polymer gels, using a 2D numerical model, in which polymer segments form a triangular network and disorder is introduced by varying their contour lengths. We show that in the relevant parameter range: (i) the non-affine contribution to the shear modulus is negligible, i.e. the Born approximation is satisfactory; (ii) the shear modulus is dominated by the contribution originating from equilibrium chain tensions. Moreover, mechanical equ…
Despite a vast clinical application of anesthetics, the molecular level of understanding of general anesthesia is far from our reach. Using atomistic molecular dynamics simulation, we study the effects of common anesthetics: ethanol, chloroform and methanol in the fully hydrated symmetric multicomponent lipid bilayer membrane comprising of an unsaturated palmitoyl-oleoyl-phosphatidyl-choline (POPC), a saturated palmitoyl-sphingomyelin (PSM) and cholesterol (Chol) which exhibits phase coexistence of liquid-ordered (lo) - liquid disordered (ld) phase domains. We find that the mechanical and phys…
Actomyosin actively generates contractile forces that provide the plasma membrane with the deformation stresses essential to carry out biological processes. Although the contractile property of purified actomyosin has been extensively studied, to understand the physical contribution of the actiomyosin contractile force on a deformable membrane is still a challenging problem and of great interest in the field of biophysics. Here, we reconstituted a model system with a cell-sized deformable interface that exhibits anomalous curvature dependent wrinkling caused by actomyosin cortex underneath the…
Spatial heterogeneity in the elastic properties of soft random solids is examined via vulcanization theory. The spatial heterogeneity in the \emph{structure} of soft random solids is a result of the fluctuations locked-in at their synthesis, which also brings heterogeneity in their \emph{elastic properties}. Vulcanization theory studies semi-microscopic models of random-solid-forming systems, and applies replica field theory to deal with their quenched disorder and thermal fluctuations. The elastic deformations of soft random solids are argued to be described by the Goldstone sector of fluctua…
An elastic membrane with embedded nematic molecules is considered as a model of anisotropic fluid membrane with internal ordering. By considering the geometric coupling between director field and membrane curvature, the nematic texture is shown to induce anisotropic stresses additional to Canham-Helfrich elasticity. Building upon differential geometry, analytical expressions are found for the membrane stress and torque induced by splaying, twisting and bending of the nematic director as described by the Frank energy of liquid crystals. The forces induced by prototypical nematic textures are vi…
The dynamics of microcapsules in steady shear flow was studied using a theoretical approach based on three variables: The Taylor deformation parameter $α_{\rm D}$, the inclination angle $θ$, and the phase angle $ϕ$ of the membrane rotation. It is found that the dynamic phase diagram shows a remarkable change with an increase in the ratio of the membrane shear and bending elasticities. A fluid vesicle (no shear elasticity) exhibits three dynamic modes: (i) Tank-treading (TT) at low viscosity $η_{\rm {in}}$ of internal fluid ($α_{\rm D}$ and $θ$ relaxes to constant values), (ii) Tumbling (TB) at…
The dynamics of drop impact on solid surfaces can be changed significantly by tuning the elasticity of the solid. Most prominently, the substrate deformation causes an increase in the splashing threshold as compared to impact onto perfectly rigid surfaces, and can thus lead to splash suppression. Here, we experimentally determine the splashing threshold for impact on thin membranes as a function of the tension in the membrane and its elastic properties. The drop dynamics is correlated to the membrane deformation, which is simultaneously measured using a laser profilometry technique. The experi…
The mechanical properties of biological membranes play an important role in the structure and the functioning of living organisms. One of the most widely used methods for determination of the bending elasticity modulus of the model lipid membranes (simplified models of the biomembranes with similar mechanical properties) is analysis of the shape fluctuations of the nearly spherical lipid vesicles. A theoretical basis of such an analysis is developed by Milner and Safran. In the present studies we analyze their results using an approach based on the Bogoljubov inequalities and the approximating…
Theoretical models describing specific adhesion of membranes predict (for certain parameters) a macroscopic phase separation of bonds into adhesion domains. We show that this behavior is fundamentally altered if the membrane is pinned randomly due to, e.g., proteins that anchor the membrane to the cytoskeleton. Perturbations which locally restrict membrane height fluctuations induce quenched disorder of the random-field type. This rigorously prevents the formation of macroscopic adhesion domains following the Imry-Ma argument [Y. Imry and S. K. Ma, Phys. Rev. Lett. 35, 1399 (1975)]. Our predic…
A nematic membrane is a sheet with embedded orientational order, which can occur in biological cells, liquid crystal films, manufactured materials, and other soft matter systems. By formulating the free energy of nematic films using tensor contractions from differential geometry, we elucidate the elastic terms allowed by symmetry, and indicate differences from hexatic membranes. We find that topological defects in the orientation field can cause the membrane to buckle over a size set by the competition between surface tension and in-plane elasticity. In the absence of bending rigidity the resu…
We show using theory and experiments that a small particle moving along an elastic membrane through a viscous fluid is repelled from the membrane due to hydro-elastic forces. The viscous stress field produces an elastic disturbance leading to particle-wave coupling. We derive an analytic expression for the particle trajectory in the lubrication limit, bypassing the construction of the detailed velocity and pressure fields. The normal force is quadratic in the parallel speed, and is a function of the tension and bending resistance of the membrane. Experimentally, we measure the normal displacem…
We present a novel approach to probe elastic properties of polyelectrolyte multilayer microcapsules. The method is based on measurements of the capsule load-deformation curves with the atomic force microscope. The experiment suggests that at low applied load deformations of the capsule shell are elastic. Using elastic theory of membranes we relate force, deformation, elastic moduli, and characteristic sizes of the capsule. Fitting to the prediction of the model yields the lower limit for Young's modulus of the polyelectrolyte multilayers of the order of 1-100 MPa, depending on the template and…
The elastic properties of a self-assembled bilayer membrane are studied using the self-consistent field theory, applied to a model system composed of flexible amphiphilic chains dissolved in hydrophilic polymeric solvents. Examining the free energy of bilayer membranes with different geometries allows us to calculate their bending modulus, Gaussian modulus, two fourth-order membrane moduli, and the line tension. The dependence of these parameters on the microscopic characteristics of the amphiphilic chain, characterized by the volume fraction of the hydrophilic component, is systematically stu…
We investigate the emergence of the continuum elastic limit from the atomistic description of matter at zero temperature considering how locally defined elastic quantities depend on the coarse graining length scale. Results obtained numerically investigating different model systems are rationalized in a unifying picture according to which the continuum elastic limit emerges through a process determined by two system properties, the degree of disorder, and a length scale associated to the transverse low-frequency vibrational modes. The degree of disorder controls the emergence of long-range loc…
We study the elasticity, fluctuations and pinning of a putative spontaneous vortex solid in ferromagnetic superconductors. Using a rigorous thermodynamic argument, we show that in the idealized case of vanishing crystalline pinning anisotropy the long-wavelength tilt modulus of such a vortex solid vanishes identically, as guaranteed by the underlying rotational invariance. The vanishing of the tilt modulus means that, to lowest order, the associated tension elasticity is replaced by the softer, curvature elasticity. The effect of this is to make the spontaneous vortex solid qualitatively more …
We examine how disordering joint position influences the linear elastic behavior of lattice materials via numerical simulations in two-dimensional beam networks. Three distinct initial crystalline geometries are selected as representative of mechanically isotropic materials low connectivity, mechanically isotropic materials with high connectivity, and mechanically anisotropic materials with intermediate connectivity. Introducing disorder generates spatial fluctuations in the elasticity tensor at the local (joint) scale. Proper coarse-graining reveals a well-defined continuum-level scale elasti…