We propose two toy-models to describe, predict, and interpret the wrinkles appearing on the surface of skin when it is sheared. With the first model, we account for the lines of greatest tension present in human skin by subjecting a layer of soft tissue to a pre-stretch, and for the epidermis by endowing one of the layer's faces with a surface tension. For the second model, we consider an anisotropic model for the skin, to reflect the presence of stiff collagen fibres in a softer elastic matrix. In both cases, we find an explicit bifurcation criterion, linking geometrical and material parameters to a critical shear deformation accompanied by small static wrinkles, with decaying amplitudes normal to the free surface of skin.
Biological processes, from morphogenesis to tumor invasion, spontaneously generate shear stresses inside living tissue. The mechanisms that govern the transmission of mechanical forces in epithelia and the collective response of the tissue to bulk shear deformations remain, however, poorly understood. Using a minimal cell-based computational model, we investigate the constitutive relation of confluent tissues under simple shear deformation. We show that an initially undeformed fluidlike tissue acquires finite rigidity above a critical applied strain. This is akin to the shear-driven rigidity o…
When a block made of an elastomer is subjected to large shear, its surface remains flat. When a block of biological soft tissue is subjected to large shear, it is likely that its surface in the plane of shear will buckle (apparition of wrinkles). One factor that distinguishes soft tissues from rubber-like solids is the presence -- sometimes visible to the naked eye -- of oriented collagen fibre bundles, which are stiffer than the elastin matrix into which they are embedded but are nonetheless flexible and extensible. Here we show that the simplest model of isotropic nonlinear elasticity, namel…
During severe impact conditions, brain tissue experiences a rapid and complex deformation, which can be seen as a mixture of compression, tension and shear. Moreover, diffuse axonal injury (DAI) occurs in animals and humans when both the strains and strain rates exceed 10% and 10/s, respectively. Knowing the mechanical properties of brain tissue in shear at these strains and strain rates is thus of particular importance, as they can be used in finite element simulations to predict the occurrence of brain injuries under different impact conditions. In this research, an experimental setup was de…
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…
We identify three distinct shearing modes for simple shear deformations of transversely isotropic soft tissue which allow for both positive and negative Poynting effects (that is, they require compressive and tensile lateral normal stresses, respectively, in order to maintain simple shear). The positive Poynting effect is that usually found for isotropic rubber. Here, specialisation of the general results to three strain-energy functions which are quadratic in the anisotropic invariants, linear in the isotropic strain invariants and consistent with the linear theory suggests that there are two…
We use supersonic shear wave imaging (SSI) technique to measure not only the linear but also the nonlinear elastic properties of brain matter. Here, we tested six porcine brains ex vivo and measured the velocities of the plane shear waves induced by acoustic radiation force at different states of pre-deformation when the ultrasonic probe is pushed into the soft tissue. We relied on an inverse method based on the theory governing the propagation of small-amplitude acoustic waves in deformed solids to interpret the experimental data. We found that, depending on the subjects, the resulting initia…
Morphogenesis depends crucially on the complex rheological properties of cell tissues and on their ability to maintain mechanical integrity while rearranging at long times. In this paper, we study the rheology of polygonal cellular networks described by a vertex model in the presence of fluctuations. We use a triangulation method to decompose shear into cell shape changes and cell rearrangements. Considering the steady-state stress under constant shear, we observe nonlinear shear-thinning behavior at all magnitudes of the fluctuations, and an even stronger nonlinear regime at lower values of t…
In biological tissues, it is now well-understood that mechanical cues are a powerful mechanism for pattern regulation. While much work has focused on interactions between cells and external substrates, recent experiments suggest that cell polarization and motility might be governed by the internal shear stiffness of nearby tissue, deemed "plithotaxis". Meanwhile, other work has demonstrated that there is a direct relationship between cell shapes and tissue shear modulus in confluent tissues. Joining these two ideas, we develop a hydrodynamic model that couples cell shape, and therefore tissue …
Shear effects in active models of normal and cancer cells
University of Texas at Austin（德克萨斯大学奥斯汀分校）；Institute for Physical Science and Technology, University of Maryland（马里兰大学物理科学与技术研究所）；Department of Chemistry, University of Texas at Austin（德克萨斯大学奥斯汀分校化学系）；Department of Physics, University of Maryland（马里兰大学物理系）；Department of Physics, University of Texas at Austin（德克萨斯大学奥斯汀分校物理系）
Rubber components and soft tissues are often subjected to large bending deformations "in service". The circumferential line elements on the inner face of a bent block can contract up to a certain critical stretch ratio lambda_cr (say) before bifurcation occurs and axial creases appear. For several models used to describe rubber, it is found that lambda_cr = 0.56, allowing for a 44% contraction. For models used to describe arteries it is found, somewhat surprisingly, that the strain-stiffening effect promotes instability. For example, the models used for the artery of a 70 year old human predic…
Mechanical signaling plays a key role in biological processes like embryo development and cancer growth. One prominent way to probe mechanical properties of tissues is to study their response to externally applied forces. Using a particle-based model featuring random apoptosis and environment-dependent division rates, we evidence a crossover from linear flow to a shear-thinning regime with increasing shear rate. To rationalize this non-linear flow we derive a theoretical mean-field scenario that accounts for the interplay of mechanical and active noise in local stresses. These noises are respe…
We experimentally observe the shear and secondary compression wave inside soft porous water-saturated melamine foams by high frame rate ultrasound imaging. Both wave speeds are supported by the weak frame of the foam. The first and second compression waves show opposite polarity, as predicted by Biot theory. Our experiments have direct implications for medical imaging: Melamine foams exhibit a similar microstructure as lung tissue. In the future, combined shear wave and slow compression wave imaging might provide new means of distinguishing malignant and healthy pulmonary tissue.
Using shear wave elastography, we measure the changes in the wave speed with the stress produced by a striated muscle during isometric voluntary contraction. To isolate the behaviour of an individual muscle from complementary or antagonistic actions of adjacent muscles, we select the flexor digiti minimi muscle, whose sole function is to extend the little finger. To link the wave speed to the stiffness, we develop an acousto-elastic theory for shear waves in homogeneous, transversely isotropic, incompressible solids subject to uniaxial stress. We then provide measurements of the apparent shear…
Adhesion differentials control the rheology of biomimetic emulsions
Sorbonne Université（索邦大学）；CNRS（法国国家科学研究中心）；Laboratoire Jean Perrin（让·佩兰实验室）；Inserm（法国国家健康与医学研究院）；Institut de Biologie Paris-Seine（巴黎塞纳生物学研究所）；Aix Marseille Univ（艾克斯-马赛大学）；Université de Toulon（土伦大学）；CPT (UMR 7332)（复杂系统物理中心（UMR 7332））；Turing Center for Living Systems（图灵生命系统中心）
