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Nonlinear stiffening is a ubiquitous property of major types of biopolymers that make up the extracellular matrices (ECM) including collagen, fibrin and basement membrane. Within the ECM, many types of cells such as fibroblasts and cancer cells are known to mechanically stretch their surroundings that locally stiffens the matrix. Although the bulk nonlinear elastic behaviors of these biopolymer networks are well studied, their local mechanical responses remain poorly characterized. Here, to understand how a living cell feels the nonlinear mechanical resistance from the ECM, we mimic the cell-applied local force using optical tweezers; we report that the local stiffening responses in highly nonlinear ECM are significantly weaker than responses found in bulk rheology, across two orders of magnitude of the locally applied force since the onset of stiffening. With a minimal model, we show that a local point force application can induce a stiffened region in the matrix, which expands with increasing magnitude of the point force. Furthermore, we show that this stiffened region behaves as an effective probe upon local loading. The local nonlinear elastic response can be attributed to the nonlinear growth of this effective probe that linearly deforms an increasing portion of the matrix.

In a diversity of physiological contexts, eukaryotic cells adhere to an extracellular matrix (ECM), a disordered network with complex nonlinear mechanics. Such cells can perform mechanosensation: using local force probing they can measure and respond to their substrate's mechanical properties. It remains unclear, however, how the mechanical complexity of the ECM at the cellular scale impacts mechanosensation. Here, we investigate the physical limits of mechanosensation imposed by the inherent structural disorder and nonlinear elastic response of the ECM. Using a theoretical framework for disordered fiber networks, we find that the extreme mechanical heterogeneity that cells can locally sense with small probing forces is strongly reduced with increasing force. Specifically, we predict that the accuracy of mechanosensation dramatically improves with force, following a universal power law insensitive to constitutive details, which we quantitatively confirm using microrheology experiments in collagen and fibrin gels. We provide conceptual insights into this behavior by introducing a general model for nonlinear mechanosensation, based on the idea of an emergent nonlinear length-scale associated with fiber buckling. This force-dependent length-scale enhances the range over which local mechanical measurements are performed, thereby averaging the response of a disordered network over an enlarged region. We show with an example how a cell can use this nonlinear mechanosensation to infer the macroscopic mechanical properties of a disordered ECM using local measurements. Together, our results demonstrate that cells can take advantage of the inherent nonlinearity of fibrous networks to robustly sense, control, and respond to their mechanical environment.

Continuum mechanics describes compressive failure as a standard bifurcation in the response of a material to an increasing load: damage, which initially grows uniformly in the material, localizes within a thin band at failure. Yet, experiments recording the acoustic activity preceding localization evidence power-law distributed failure precursors of increasing size, suggesting that compressive failure is a critical phenomenon. We examine here this apparent contradiction by probing the spatial organization of the damage activity and its evolution until localization during compression experiments of 2D cellular solids. The intermittent damage evolution measured in our experiments is adequately described by a non-stationary depinning equation derived from damage mechanics and reminiscent of critical phenomena. In this description, precursors are damage cascades emerging from the interplay between the material's disorder and the long-range stress redistributions following individual damage events. Yet, the divergence of their characteristic size close to failure, which we observe in our experiments, is not the signature of a transition towards criticality. Instead, the system remains at a fixed distance to the critical point at all stages of the damage evolution. The divergence results from the progressive loss of stability of the material as it is driven towards localization. Thus, our study shows that compressive failure is a standard bifurcation for which the material disorder plays a marginal role. It also shows that precursory activity constitute by-products of the evolution towards localization and can serve to build a predictive method to assess the residual lifetime of structures., Comment: 35 pages, 14 figures including the supplementary information

Forecasting fracture locations in a progressively failing disordered structure is of paramount importance when considering structural materials. We explore this issue for gradual deterioration via beam breakage of two-dimensional disordered lattices, which we represent as networks, for various values of mean degree. We study experimental samples with geometric structures that we construct based on observed contact networks in 2D granular media. We calculate geodesic edge betweenness centrality, which helps quantify which edges are on many shortest paths in a network, to forecast the failure locations. We demonstrate for the tested samples that, for a variety of failure behaviors, failures occur predominantly at locations that have larger geodesic edge betweenness values than the mean one in the structure. Because only a small fraction of edges have values above the mean, this is a relevant diagnostic to assess failure locations. Our results demonstrate that one can consider only specific parts of a system as likely failure locations and that, with reasonable success, one can assess possible failure locations of a structure without needing to study its detailed energetic states., Comment: Extended discussion, added supplementary information

In ordinary solids, material disorder is known to increase the size of the process zone in which stress concentrates at the crack tip, causing a transition from localized to diffuse failure. Here, we report experiments on disordered 2D lattices, derived from frictional particle packings, in which the mean coordination number $\langle z \rangle$ of the underlying network provides a similar control. Our experiments show that tuning the connectivity of the network provides access to a range of behaviors from brittle to ductile failure. We elucidate the cooperative origins of this transition using a frictional pebble game algorithm on the original, intact lattices. We find that the transition corresponds to the isostatic value $\langle z \rangle = 3$ in the large-friction limit, with brittle failure occurring for structures vertically spanned by a rigid cluster, and ductile failure for floppy networks containing nonspanning rigid clusters. Furthermore, we find that individual failure events typically occur within the floppy regions separated by the rigid clusters.

The failure of brittle solids involves, before macroscopic rupture, power-law distributed avalanches of local rupture events whereby microcracks nucleate and grow, which are also observed in for an elastic interface evolving in a non-homogeneous medium. For this reason, it is tempting to relate failure to the depinning of an elastic interface. Here we compute the elastic kernel of the interface representing the damage field of a brittle solid. In the case of a damage model of rupture under compression, which implements the Mohr-Coulomb criterion at the local scale, we show that the elastic kernel is unstable, and hence is very different from the kernels of usual interfaces. We show that the unstable modes are responsible for the localization of damage along a macroscopic fault observed in numerical simulations. At low disorder, the most unstable mode gives the orientation of the macroscopic fault that we measure in numerical simulations. The orientation of the fault changes when the level of disorder is increased, suggesting a complex interplay of the unstable modes and the disorder.

The Mohr-Coulomb criterion is widely used in geosciences to relate the state of stress at failure to the observed orientation of the resulting faults. This relation is based on the assumption that the fault occurs along a plane that maximizes the Coulomb stress. Here, we test this hypothesis using an elastic, progressive damage model that implements the Mohr-Coulomb criterion at the local scale. We find that the orientation of the fault is not given by the Mohr-Coulomb criterion. Instead, for minimal disorder, it corresponds to the most unstable mode of damage in the model, which we determine through a linear stability analysis of the homogeneously damaged state. Our simulations show that microstructural disorder significantly affects the orientation of the fault, which, however, remains always far from the Mohr-Coulomb prediction.