A computational framework for the morpho-elastic development of molluskan shells by surface and volume growth

Mollusk shells are an ideal model system for understanding the morpho-elastic basis of morphological evolution of invertebrates' exoskeletons. During the formation of the shell, the mantle tissue secretes proteins and minerals that calcify to form a new incremental layer of the exoskeleton. Most of the existing literature on the morphology of mollusks is descriptive. The mathematical understanding of the underlying coupling between pre-existing shell morphology, de novo surface deposition and morpho-elastic volume growth is at a nascent stage, primarily limited to reduced geometric representations. Here, we propose a general, three-dimensional computational framework coupling pre-existing morphology, incremental surface growth by accretion, and morpho-elastic volume growth. We exercise this framework by applying it to explain the stepwise morphogenesis of seashells during growth: new material surfaces are laid down by accretive growth on the mantle whose form is determined by its morpho-elastic growth. Calcification of the newest surfaces extends the shell as well as creates a new scaffold that constrains the next growth step. We study the effects of surface and volumetric growth rates, and of previously deposited shell geometries on the resulting modes of mantle deformation, and therefore of the developing shell's morphology. Connections are made to a range of complex shells ornamentations.Comment: Main article is 20 pages long with 15 figures. Supplementary material is 4 pages long with 6 figures and 6 attached movies. To be published in PLOS Computational Biolog

Morphoelastic rods Part 1: A single growing elastic rod

A theory for the dynamics and statics of growing elastic rods is presented. First, a single growing rod is considered and the formalism of three-dimensional multiplicative decomposition of morphoelasticity is used to describe the bulk growth of Kirchhoff elastic rods. Possible constitutive laws for growth are discussed and analysed. Second, a rod constrained or glued to a rigid substrate is considered, with the mismatch between the attachment site and the growing rod inducing stress. This stress can eventually lead to instability, bifurcation, and buckling

Indentation of ellipsoidal and cylindrical elastic shells

Thin shells are found in nature at scales ranging from viruses to hens’ eggs; the stiffness of such shells is essential for their function. We present the results of numerical simulations and theoretical analyses for the indentation of ellipsoidal and cylindrical elastic shells, considering both pressurized and unpressurized shells. We provide a theoretical foundation for the experimental findings of Lazarus et al. [Phys. Rev. Lett. (submitted)] and for previous work inferring the turgor pressure of bacteria from measurements of their indentation stiffness; we also identify a new regime at large indentation. We show that the indentation stiffness of convex shells is dominated by either the mean or Gaussian curvature of the shell depending on the pressurization and indentation depth. Our results reveal how geometry rules the rigidity of shells

Nonlinear morphoelastic plates II: exodus to buckled states

Morphoelasticity is the theory of growing elastic materials. This theory is based on the multiple decomposition of the deformation gradient and provides a formulation of the deformation and stresses induced by growth. Following a companion paper, a general theory of growing nonlinear elastic Kirchhoff plate is described. First, a complete geometric description of incompatibility with simple examples is given. Second, the stability of growing Kirchhoff plates is analyzed

Preconditioning for Allen-Cahn variational inequalities with non-local constraints

The solution of Allen-Cahn variational inequalities with mass constraints is of interest in many applications. This problem can be solved both in its scalar and vector-valued form as a PDE-constrained optimization problem by means of a primal-dual active set method. At the heart of this method lies the solution of linear systems in saddle point form. In this paper we propose the use of Krylov-subspace solvers and suitable preconditioners for the saddle point systems. Numerical results illustrate the competitiveness of this approach

Morphing of Geometric Composites via Residual Swelling

Understanding and controlling the shape of thin, soft objects has been the focus of significant research efforts among physicists, biologists, and engineers in the last decade. These studies aim to utilize advanced materials in novel, adaptive ways such as fabricating smart actuators or mimicking living tissues. Here, we present the controlled growth--like morphing of 2D sheets into 3D shapes by preparing geometric composite structures that deform by residual swelling. The morphing of these geometric composites is dictated by both swelling and geometry, with diffusion controlling the swelling-induced actuation, and geometric confinement dictating the structure's deformed shape. Building on a simple mechanical analog, we present an analytical model that quantitatively describes how the Gaussian and mean curvatures of a thin disk are affected by the interplay among geometry, mechanics, and swelling. This model is in excellent agreement with our experiments and numerics. We show that the dynamics of residual swelling is dictated by a competition between two characteristic diffusive length scales governed by geometry. Our results provide the first 2D analog of Timoshenko's classical formula for the thermal bending of bimetallic beams - our generalization explains how the Gaussian curvature of a 2D geometric composite is affected by geometry and elasticity. The understanding conferred by these results suggests that the controlled shaping of geometric composites may provide a simple complement to traditional manufacturing techniques

Circumferential buckling instability of a growing cylindrical tube

A cylindrical elastic tube under uniform radial pressure will buckle circumferentially to a non-circular cross section at a critical pressure. The buckling represents an instability of the inner or outer edge of the tube. This is a common phenomenon in biological tissues, where it is referred to as mucosal folding. Here, we investigate this buckling instability in a growing elastic tube. A change in thickness due to growth can have a dramatic impact on circumferential buckling, both in the critical pressure and the buckling pattern. We consider both single and bi-layer tubes and multiple boundary conditions. We highlight the competition between geometric effects, i.e. the change in tube dimensions, and mechanical effects, i.e. the effect of residual stress, due to differential growth. This competition can lead to non-intuitive results, such as a tube growing to be thicker and yet buckle at a lower pressure