Collagen is the most abundant extracellular-network-forming protein in animal biology and is important in both natural and artificial tissues, where it serves as a material of great mechanical versatility. This versatility arises from its almost unique ability to remodel under applied loads into anisotropic and inhomogeneous structures. To explore the origins of this property, we develop a set of analysis tools and a novel experimental setup that probes the mechanical response of fibrous networks in a geometry that mimics a typical deformation profile imposed by cells in vivo. We observe strong fiber alignment and densification as a function of applied strain for both uncrosslinked and crosslinked collagenous networks. This alignment is found to be irreversibly imprinted in uncrosslinked collagen networks, suggesting a simple mechanism for tissue organization at the microscale. However, crosslinked networks display similar fiber alignment and the same geometrical properties as uncrosslinked gels, but with full reversibility. Plasticity is therefore not required to align fibers. On the contrary, our data show that this effect is part of the fundamental non-linear properties of fibrous biological networks.
Collagen is the most abundant extracellular-matrix protein in mammals and the main structural and load-bearing element of connective tissues. Collagen networks show remarkable strain-stiffening properties which tune the mechanical functions of tissues and regulate cell behaviours. Linear and non-linear mechanical properties of in-vitro disordered collagen networks have been widely studied using rheology for a range of self-assembly conditions in recent years. However, a one to one correlation between the onset of macroscopic network failure and local deformations inside the sample is yet to be…
Collagen fibrils are the main structural component of load-bearing tissues such as tendons, ligaments, skin, the cornea of the eye, and the heart. The D-band of collagen fibrils is an axial periodic density modulation that can be easily characterized by tissue-level X-ray scattering. During mechanical testing, D-band strain is often used as a proxy for fibril strain. However, this approach ignores the coupling between strain and molecular tilt. We examine the validity of this approximation using an elastomeric collagen fibril model that includes both the D-band and a molecular tilt field. In t…
Collagen forms the structural scaffold of connective tissues in all mammals. Tissues are remarkably resistant against mechanical deformations because collagen molecules hierarchically self-assemble in fibrous networks that stiffen with increasing strain. Nevertheless, collagen networks do fracture when tissues are overloaded or subject to pathological conditions such as aneurysms. Prior studies of the role of collagen in tissue fracture have mainly focused on tendons, which contain highly aligned bundles of collagen. By contrast, little is known about fracture of the orientationally more disor…
Protein-stabilised emulsion gels can be studied in the theoretical framework of colloidal gels, because both protein assemblies and droplets may be considered as soft colloids. These particles differ in their nature, size and softness, and these differences may have an influence on the rheological properties of the gels they form. Pure gels made of milk proteins (sodium caseinate), or of sub-micron protein-stabilised droplets, were prepared by slow acidification of suspensions at various concentrations. Their microstructure was characterised, their viscoelasticity, both in the linear and non-l…
Living tissues show an extraordinary adaptiveness to strain, which is crucial for their proper biological functioning. The physical origin of this mechanical behaviour has been widely investigated using reconstituted networks of collagen fibres, the principal load-bearing component of tissues. However, collagen fibres in tissues are embedded in a soft hydrated polysaccharide matrix which generates substantial internal stresses whose effect on tissue mechanics is unknown. Here, by combining mechanical measurements and computer simulations, we show that networks composed of collagen fibres and a…
Stress induced yielding/fluidization in disordered solids, characterized by irreversibility and enhanced dissipation, is important for a wide range of industrial and geological processes. Although, such phenomena in thermal systems have been extensively studied, they remain poorly understood for granular solids. Here, using oscillatory shear rheology and in-situ optical imaging, we study energy dissipation in a dense granular suspension of adhesive particles that forms yield stress solids far below the isotropic jamming point obtained in the limit of hard-sphere repulsion. We find interesting …
Biopolymer networks are common in biological systems from the cytoskeleton of individual cells to collagen in the extracellular matrix. The mechanics of these systems under applied strain can be explained in some cases by a phase transition from soft to rigid states. For collagen networks, it has been shown that this transition is critical in nature and it is predicted to exhibit diverging fluctuations near a critical strain that depends on the network's connectivity and structure. Whereas prior work focused mostly on shear deformation that is more accessible experimentally, here we study the …
Understanding the origin of the unique rheological properties of wheat gluten, the protein fraction of wheat grain, is crucial in bread-making processes and questions scientists since polymeric glutenins. To better understand the respective role of the different classes of proteins in the supramolecular structure of gluten and its link to the material properties, we investigate here concentrated dispersions of gluten proteins in water with a fixed total protein concentration but variable composition in gliadin and glutenin. Linear viscoelasticity measurements show a gradual increase of the vis…
Unlike hard materials such as metals and ceramics, rubbery materials can endure large deformations due to the large conformational degree of freedom of the crosslinked three-dimensional polymer network. However, the effect of the branching factor of the network on the ultimate mechanical properties of rubbery materials has not yet been clarified. This study shows that tri-branching, which entails the lowest branching factor, results in a large elastic deformation near the theoretical upper bound. This ideal elastic limit is realized by reversible strain-induced crystallization, providing on-de…
We present a detailed numerical study of multi-component colloidal gels interacting sterically and obtained by arrested phase separation. Under deformation, we found that the interplay between the different intertwined networks is key. Increasing the number of component leads to softer solids that can accomodate progressively larger strain before yielding. The simulations highlight how this is the direct consequence of the purely repulsive interactions between the different components, which end up enhancing the linear response of the material. Our work {provides new insight into mechanisms at…
We present a Landau type theory for the non-linear elasticity of biopolymer gels with a part of the order parameter describing induced nematic order of fibers in the gel. We attribute the non-linear elastic behavior of these materials to fiber alignment induced by strain. We suggest an application to contact guidance of cell motility in tissue. We compare our theory to simulation of a disordered lattice model for biopolymers. We treat homogeneous deformations such as simple shear, hydrostatic expansion, and simple extension, and obtain good agreement between theory and simulation. We also cons…
We propose a theory based on non-equilibrium thermodynamics to describe the mechanical behavior of an active polymer gel created by the inclusion of molecular motors in its solvent. When activated, these motors attach to the chains of the polymer network and shorten them creating a global contraction of the gel, which mimics the active behavior of a cytoskeleton. The power generated by these motors is obtained by ATP hydrolysis reaction, which transduces chemical energy into mechanical work. The latter is described by an increment of strain energy in the gel due to an increased stiffness. This…
Colloidal gels made of carbon black particles dispersed in light mineral oil are "rheo-acoustic" materials, i.e., their mechanical and structural properties can be tuned using high-power ultrasound, sound waves with submicron amplitude and frequency larger than 20~kHz . The effects of high-power ultrasound on the carbon black gel are demonstrated using two experiments: rheology coupled to ultrasound to test for the gel mechanical response and a timeresolved ultra small-angle X-ray scattering experiment (TRUSAXS) coupled to ultrasound to test for structural changes within the gel. We show that …
Mucus is a biological gel covering the surface of several tissues and insuring key biological functions, including as a protective barrier against dehydration, pathogens penetration, or gastric acids. Mucus biological functioning requires a finely tuned balance between solid-like and fluid-like mechanical response, insured by reversible bonds between mucins, the glycoproteins that form the gel. In living organisms, mucus is subject to various kinds of mechanical stresses, e.g. due to osmosis, bacterial penetration, coughing and gastric peristalsis. However, our knowledge of the effects of stre…
Filamentous bio-materials such as fibrin or collagen networks exhibit an enormous stiffening of their elastic moduli upon large deformations. This pronounced nonlinear behavior stems from a significant separation between the stiffnesses scales associated with bending vs. stretching the material's constituent elements. Here we study a simple model of such materials - floppy networks of hinged rigid bars embedded in an elastic matrix - in which the effective ratio of bending to stretching stiffnesses vanishes identically. We introduce a theoretical framework and build upon it to construct a nume…
Physically assembled gels have promising applications in many fields because of their tunable mechanical properties. Here, we report the mechanical properties as a function of polymer volume fraction ($ϕ$) for a physical gel system consists of poly(styrene)-poly(isoprene)-poly(styrene) [PS-PI-PS] in mineral oil. The PI-block molecular weight is higher than the entanglement molecular weight, which leads to the entanglement of PI-blocks at higher $ϕ$. The micellar microstructure for all gels results in a similar stress relaxation mechanism, as captured by the superposition of stress-relaxation r…
Brain tissue is a heterogeneous material, constituted by a soft matrix filled with cerebrospinal fluid. The interactions between, and the complexity of each of these components are responsible for the non-linear rate-dependent behaviour that characterizes what is one of the most complex tissue in nature. Here, we investigate the influence of the cutting rate on the fracture properties of brain, through wire cutting experiments. We also present a model for the rate-dependent behaviour of fracture propagation in soft materials, which comprises the effects of fluid interaction through a poro-hype…
Through mechanical forces, biological cells remodel the surrounding collagen network, generating striking deformation patterns. Tethers-tracts of high densification and fiber alignment-form between cells, thinner bands emanate from cell clusters. While tethers facilitate cell migration and communication, how they form is unclear. Combining modeling, simulation and experiment, we show that tether formation is a densification phase transition of the extracellular matrix, caused by buckling instability of network fibers under cell-induced compression, featuring unexpected similarities with marten…
