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We derive and simulate a mathematical model for mechanotransduction related to the Rho GTPase signalling pathway. The model addresses the bidirectional coupling between signalling processes and cell mechanics. A numerical method based on bulk-surface finite elements is proposed for the approximation of the coupled system of nonlinear reaction-diffusion equations, defined inside the cell and on the cell membrane, and the equations of elasticity. Our simulation results illustrate novel emergent features such as the strong dependence of the dynamics on cell shape, a threshold-like response to changes in substrate stiffness, and the fact that coupling mechanics and signalling can lead to the robustness of cell deformation to larger changes in substrate stiffness, ensuring mechanical homeostasis in agreement with experiments., Comment: Code available on request

We consider the numerical approximation of the stochastic complex Ginzburg-Landau equation with additive noise on the one dimensional torus. The complex nature of the equation means that many of the standard approaches developed for stochastic partial differential equations can not be directly applied. We use an energy approach to prove an existence and uniqueness result as well to obtain moment bounds on the stochastic PDE before introducing our numerical discretization. For such a well studied deterministic equation it is perhaps surprising that its numerical approximation in the stochastic setting has not been considered before. Our method is based on a spectral discretization in space and a Lie-Trotter splitting method in time. We obtain moment bounds for the numerical method before proving our main result: strong convergence on a set of arbitrarily large probability. From this we obtain a result on convergence in probability. We conclude with some numerical experiments that illustrate the effectiveness of our method., Comment: 22 pages, 8 figures

In this paper we consider the multiscale modelling of water transport in vegetated soil. In the microscopic model we distinguish between subdomains of soil and plant tissue, and use the Richards equation to model the water transport through each. Water uptake is incorporated by means of a boundary condition on the surface between root tissue and soil. Assuming a simplified root system architecture, which gives a cylindrical microstructure to the domain, the two-scale convergence and periodic unfolding methods are applied to rigorously derive a macroscopic model for water transport in vegetated soil. The degeneracy of the Richards equation and the dependence of root tissue permeability on the small parameter introduce considerable challenges in deriving macroscopic equations, especially in proving strong convergence. The variable-doubling method is used to prove the uniqueness of solutions to the model, and also to show strong two-scale convergence in the non-linear terms of the equation for water transport through root tissue.

We investigate the effect of a Heaviside cut-off on the front propagation dynamics of the so-called Burgers-FisherKolmogoroff-Petrowskii-Piscounov (Burgers-FKPP) advection-reaction-diffusion equation. We prove the existence and uniqueness of a travelling front solution in the presence of a cut-off in the reaction kinetics and the advection term, and we derive the leading-order asymptotics for the speed of propagation of the front in dependence on the advection strength and the cut-off parameter. Our analysis relies on geometric techniques from dynamical systems theory and specifically, on geometric desingularisation, which also known as blow-up.

In this paper, we consider effective discretization strategies and iterative solvers for nonlinear PDE-constrained optimization models for pattern evolution within biological processes. Upon a Sequential Quadratic Programming linearization of the optimization problem, we devise appropriate time-stepping schemes and discrete approximations of the cost functionals such that the discretization and optimization operations are commutative, a highly desirable property of a discretization of such problems. We formulate the large-scale, coupled linear systems in such a way that efficient preconditioned iterative methods can be applied within a Krylov subspace solver. Numerical experiments demonstrate the viability and efficiency of our approach., Comment: 33 pages, 3 figures, 7 tables

Collective cell migration is a multicellular phenomenon that arises in various biological contexts, including cancer and embryo development. "Collectiveness" can be promoted by cell-cell interactions such as co-attraction and contact inhibition of locomotion. These mechanisms act on cell polarity, pivotal for directed cell motility, through influencing the intracellular dynamics of small GTPases such as Rac1. To model these dynamics we introduce a biased random walk model, where the bias depends on the internal state of Rac1, and the Rac1 state is influenced by cell-cell interactions and chemoattractive cues. In an extensive simulation study we demonstrate and explain the scope and applicability of the introduced model in various scenarios. The use of a biased random walk model allows for the derivation of a corresponding partial differential equation for the cell density while still maintaining a certain level of intracellular detail from the individual based setting.

How morphogenesis depends on cell properties is an active direction of research. Here, we focus on mechanical models of growing plant tissues, where microscopic (sub)cellular structure is taken into account. In order to establish links between microscopic and macroscopic tissue properties, we perform a multiscale analysis of a model of growing plant tissue with subcellular resolution. We use homogenization to rigorously derive the corresponding macroscopic tissue scale model. Tissue scale mechanical properties are computed from microscopic structural and material properties, taking into account deformation by the growth field. We then consider case studies and numerically compare the detailed microscopic model and the tissue-scale model, both implemented using finite element method. We find that the macroscopic model can be used to efficiently make predictions about several configurations of interest. Our work will help making links between microscopic measurements and macroscopic observations in growing tissues.

We develop a numerical method for the Westervelt equation, an important equation in nonlinear acoustics, in the form where the attenuation is represented by a class of non-local in time operators. A semi-discretisation in time based on the trapezoidal rule and A-stable convolution quadrature is stated and analysed. Existence and regularity analysis of the continuous equations informs the stability and error analysis of the semi-discrete system. The error analysis includes the consideration of the singularity at $t = 0$ which is addressed by the use of a correction in the numerical scheme. Extensive numerical experiments confirm the theory.

We study a singularly perturbed fast-slow system of two partial differential equations (PDEs) of reaction-diffusion type on a bounded domain via Galerkin discretisation. We assume that the reaction kinetics in the fast variable realise a generic fold singularity, whereas the slow variable takes the role of a dynamic bifurcation parameter, thus extending the classical analysis of the singularly perturbed fold. Our approach combines a spectral Galerkin discretisation with techniques from geometric singular perturbation theory which are applied to the resulting high-dimensional systems of ordinary differential equations. In particular, we show the existence of invariant slow manifolds in the phase space of the original system of PDEs away from the fold singularity, while the passage past the singularity of the Galerkin manifolds obtained after discretisation is described by geometric desingularisation, or blow-up. Finally, we discuss the relation between these Galerkin manifolds and the underlying slow manifolds., Comment: 45 pages with 9 figures. Corresponding author: Thomas Zacharis

In this article a fractional cross-diffusion system is derived as the rigorous many-particle limit of a multi-species system of moderately interacting particles that is driven by L\'{e}vy noise. The form of the mutual interaction is motivated by the porous medium equation with fractional potential pressure. Our approach is based on the techniques developed by Oelschl\"ager (1989) and Stevens (2000), in the latter of which the convergence of a regularization of the empirical measure to the solution of a correspondingly regularized macroscopic system is shown. A well-posedness result and the non-negativity of solutions are proved for the regularized macroscopic system, which then yields the same results for the non-regularized fractional cross-diffusion system in the limit.

In this paper we undertake a multiscale analysis of nutrient uptake by plant root hairs, considering different scale relations between the radius of hairs and the distance between them. We combine the method of formal asymptotic expansions and rigorous derivation of macroscopic equations. The former prompt us to study a distinguished limit (which yields a distinct effective equation), allow us to determine higher order correctors and provide motivation for the construction of correctors essential for rigorous derivation of macroscopic equations. In the final section, we validate the results of our asymptotic analysis by direct comparison with full-geometry numerical simulations., Comment: 30 pages, 6 figures, accepted for publication in SIAM Journal on Applied Mathematics

The abstract chemical reaction $$ A+A \to \emptyset, $$ understood as a Markov chain in continuous time, has been studied in the physical literature for several years. It has been claimed that this reaction can be described by means of the stochastic differential equation $$ d \phi = - \phi^2 dt + i \, \phi \, dW_t, $$ where $i$ is the imaginary unit. This affirmation is, at least, intriguing, and has led to controversy and criticisms in the literature. The goal of this work is to give partial evidence that such a description may be possible.

In this paper homogenization of a mathematical model for biomechanics of a plant tissue with randomly distributed cells is considered. Mechanical properties of a plant tissue are modelled by a strongly coupled system of reaction-diffusion-convection equations for chemical processes in plant cells and cell walls, the equations of poroelasticity for elastic deformations of plant cell walls and middle lamella, and the Stokes equations for fluid flow inside the cells. The nonlinear coupling between the mechanics and chemistry is given by the dependence of elastic properties of plant tissue on densities of chemical substances as well as by the dependence of chemical reactions on mechanical stresses present in a tissue. Using techniques of stochastic homogenization we derive rigorously macroscopic model for plant tissue biomechanics with random distribution of cells. Strong stochastic two-scale convergence is shown to pass to the limit in the non-linear reaction terms. Appropriate meaning of the boundary terms is introduced to define the macroscopic equations with flux boundary conditions and transmission conditions on the microscopic scale.

In this paper we present an individual-based mechanical model that describes the dynamics of two contiguous cell populations with different proliferative and mechanical characteristics. An off-lattice modelling approach is considered whereby: (i) every cell is identified by the position of its centre; (ii) mechanical interactions between cells are described via generic nonlinear force laws; and (iii) cell proliferation is contact inhibited. We formally show that the continuum counterpart of this discrete model is given by a free-boundary problem for the cell densities. The results of the derivation demonstrate how the parameters of continuum mechanical models of multicellular systems can be related to biophysical cell properties. We prove an existence result for the free-boundary problem and construct travelling-wave solutions. Numerical simulations are performed in the case where the cellular interaction forces are described by the celebrated Johnson-Kendall-Roberts model of elastic contact, which has been previously used to model cell-cell interactions. The results obtained indicate excellent agreement between the simulation results for the individual-based model, the numerical solutions of the corresponding free-boundary problem and the travelling-wave analysis.

Plant hormone auxin has critical roles in plant growth, dependent on its heterogeneous distribution in plant tissues. Exactly how auxin transport and developmental processes such as growth coordinate to achieve the precise patterns of auxin observed experimentally is not well understood. Here we use mathematical modelling to examine the interplay between auxin dynamics and growth and their contribution to formation of patterns in auxin distribution in plant tissues. Mathematical models describing the auxin-related signalling pathway, PIN and AUX1 dynamics, auxin transport, and cell growth in plant tissues are derived. A key assumption of our models is the regulation of PIN proteins by the auxin-responsive ARF-Aux/IAA signalling pathway, with upregulation of PIN biosynthesis by ARFs. Models are analysed and solved numerically to examine the long-time behaviour and auxin distribution. Changes in auxin-related signalling processes are shown to be able to trigger transition between passage and spot type patterns in auxin distribution. The model was also shown to be able to generate isolated cells with oscillatory dynamics in levels of components of the auxin signalling pathway which could explain oscillations in levels of ARF targets that have been observed experimentally. Cell growth was shown to have influence on PIN polarisation and determination of auxin distribution patterns. Numerical simulation results indicate that auxin-related signalling processes can explain the different patterns in auxin distributions observed in plant tissues, whereas the interplay between auxin transport and growth can explain the `reverse-fountain' pattern in auxin distribution observed at plant root tips.

Two-scale homogenization limits of parabolic cross-diffusion systems in a heterogeneous medium with no-flux boundary conditions are proved. The heterogeneity of the medium is reflected in the diffusion coefficients or by the perforated domain. The diffusion matrix is of degenerate type and may be neither symmetric nor positive semi-definite, but the diffusion system is assumed to satisfy an entropy structure. Uniform estimates are derived from the entropy production inequality. New estimates on the equicontinuity with respect to the time variable ensure the strong convergence of a sequence of solutions to the microscopic problems defined in perforated domains.

The Alber equation is a moment equation for the nonlinear Schr\"odinger equation, formally used in ocean engineering to investigate the stability of stationary and homogeneous sea states in terms of their power spectra. In this work we present the first well-posedness theory for the Alber equation with the help of an appropriate equivalent reformulation. Moreover, we show linear Landau damping in the sense that, under a stability condition on the homogeneous background, any inhomogeneities disperse and decay in time. The proof exploits novel $L^2$ space-time estimates to control the inhomogeneity and our result applies to any regular initial data (without a mean-zero restriction). Finally, the sufficient condition for stability is resolved, and the physical implications for ocean waves are discussed. Using a standard reference dataset (the "North Atlantic Scatter Diagram") it is found that the vast majority of sea states are stable, but modulationally unstable sea states do appear, with likelihood $O(1/1000);$ these would be the prime breeding ground for rogue waves.

We present and analyse a model for cell signalling processes in biological tissues. The model includes diffusion and nonlinear reactions on the cell surfaces, and both inter- and intracellular signalling. Using techniques from the theory of two-scale convergence as well the unfolding method, we show convergence of the solutions to the model to solutions of a two-scale macroscopic problem. We also present a two-scale bulk-surface finite element method for the approximation of the macroscopic model. We report on some benchmarking results as well as numerical simulations in a biologically relevant regime that illustrate the influence of cell-scale heterogeneities on macroscopic concentrations., Comment: Accepted for publication in SIAM MMS

Multiscale analysis of a degenerate pseudoparabolic variational inequality, modelling the two-phase flow with dynamical capillary pressure in a perforated domain, is the main topic of this work. Regularisation and penalty operator methods are applied to show the existence of a solution of the nonlinear degenerate pseudoparabolic variational inequality defined in a domain with microscopic perforations, as well as to derive a priori estimates for solutions of the microscopic problem. The main challenge is the derivation of a priori estimates for solutions of the variational inequality, uniformly with respect to the regularisation parameter and to the small parameter defining the scale of the microstructure. The method of two-scale convergence is used to derive the corresponding macroscopic obstacle problem.

The plant hormones brassinosteroid (BR) and gibberellin (GA) have important roles in a wide range of processes involved in plant growth and development. In this paper we derive and analyse new mathematical models for the BR signalling pathway and for the crosstalk between the BR and GA signalling pathways. To analyse the effects of spatial heterogeneity of the signalling processes, along with spatially-homogeneous ODE models we derive coupled PDE-ODE systems modelling the temporal and spatial dynamics of molecules involved in the signalling pathways. The values of the parameters in the model for the BR signalling pathway are determined using experimental data on the gene expression of BR biosynthetic enzymes. The stability of steady state solutions of our mathematical model, shown for a wide range of parameters, can be related to the BR homeostasis. Our results for the crosstalk model suggest that the interaction between transcription factors BZR and DELLA exerts more influence on the dynamics of the signalling pathways than BZR-mediated biosynthesis of GA, suggesting that the interaction between transcription factors may constitute the principal mechanism of the crosstalk between the BR and GA signalling pathways. In general, perturbations in the GA signalling pathway have larger effects on the dynamics of components of the BR signalling pathway than perturbations in the BR signalling pathway on the dynamics of the GA pathway. The perturbation in the crosstalk mechanism also has a larger effect on the dynamics of the BR pathway than of the GA pathway. Large changes in the dynamics of the GA signalling processes can be observed only in the case where there are disturbances in both the BR signalling pathway and the crosstalk mechanism., Comment: Journal of Theoretical Biology 2017

In this paper a microscopic model for a signalling process in the cardiac muscle tissue of the left ventricular wall, comprising non-periodic fibrous microstructure is considered. To derive the macroscopic equations we approximate the non-periodic microstructure by the corresponding locally-periodic microstructure. Then applying the methods of the locally-periodic (l-p) unfolding operator, locally-periodic two-scale (l-t-s) convergence on oscillating surfaces and l-p boundary unfolding operator we obtain the macroscopic problem for a signalling process in the heart muscle tissue.

The microscopic structure of a plant cell wall is given by cellulose microfibrils embedded in a cell wall matrix. In this paper we consider a microscopic model for interactions between viscoelastic deformations of a plant cell wall and chemical processes in the cell wall matrix. We consider elastic deformations of the cell wall microfibrils and viscoelastic Kelvin--Voigt type deformations of the cell wall matrix. Using homogenization techniques (two-scale convergence and periodic unfolding methods) we derive macroscopic equations from the microscopic model for cell wall biomechanics consisting of strongly coupled equations of linear viscoelasticity and a system of reaction-diffusion and ordinary differential equations. As is typical for microscopic viscoelastic problems, the macroscopic equations for viscoelastic deformations of plant cell walls contain memory terms. The derivation of the macroscopic problem for degenerate viscoelastic equations is conducted using a perturbation argument., Comment: arXiv admin note: substantial text overlap with arXiv:1410.6911

It is known that the orientation of cellulose microfibrils within plant cell walls has an important impact on the morphogenesis of plant cells and tissues. Viewing the shape of a plant cell as a square prism or cylinder with the axis aligning with the primary direction of expansion and growth, the orientation of the microfibrils within the cell wall on the sides of the cell is known. However, not much is known about their orientation at the ends of the cell. Here we investigate the impact of the orientation of cellulose microfibrils within a plant cell wall at the ends of the cell by solving the equations of linear elasticity numerically. Three different scenarios for the orientation of the microfibrils are considered. The macroscopic elastic properties of the cell wall are obtained using homogenization theory from the microscopic description of the elastic properties of the cell wall microfibrils and wall matrix. It is found that the orientation of the microfibrils in the upper and lower parts of cell walls do not affect the expansion of the cell in the direction of its axis but do affect its expansion in the lateral directions. The arrangement of the microfibrils in the upper and lower parts of cell walls is especially important in the case of directed forces acting on plant cell walls and tissues., Comment: 16 pages, 7 figures

In this paper homogenization of a mathematical model for plant tissue biomechanics is presented. The microscopic model constitutes a strongly coupled system of reaction-diffusion-convection equations for chemical processes in plant cells, the equations of poroelasticity for elastic deformations of plant cell walls and middle lamella, and Stokes equations for fluid flow inside the cells. The chemical process in cells and the elastic properties of cell walls and middle lamella are coupled because elastic moduli depend on densities involved in chemical reactions, whereas chemical reactions depend on mechanical stresses. Using homogenization techniques we derive rigorously a macroscopic model for plant biomechanics. To pass to the limit in the nonlinear reaction terms, which depend on elastic strain, we prove the strong two-scale convergence of the displacement gradient and velocity field.

Here homogenization theory is used to establish a connection between the symmetries of a periodic elastic structure associated with the microscopic properties of an elastic material and the material symmetries of the effective, macroscopic elasticity tensor. Previous results of this type exist but here more general symmetries on the microscale are considered. Using an explicit example, we show that it is possible for a material to be fully anisotropic on the microscale and yet the symmetry group on the macroscale can contain elements other than plus or minus the identity. Another example demon- strates that not all material symmetries of the macroscopic elastic tensor are generated by symmetries of the periodic elastic structure., Comment: 18 pages, 5 figures

In this paper we present a derivation and multiscale analysis of a mathematical model for plant cell wall biomechanics that takes into account both the microscopic structure of a cell wall coming from the cellulose microfibrils and the chemical reactions between the cell wall's constituents. Particular attention is paid to the role of pectin and the impact of calcium-pectin cross-linking chemistry on the mechanical properties of the cell wall. We prove the existence and uniqueness of the strongly coupled microscopic problem consisting of the equations of linear elasticity and a system of reaction-diffusion and ordinary differential equations. Using homogenization techniques (two-scale convergence and periodic unfolding methods) we derive a macroscopic model for plant cell wall biomechanics.

In this paper we generalize the periodic unfolding method and the notion of two-scale convergence on surfaces of periodic microstructures to locally periodic situations. The methods that we introduce allow us to consider a wide range of non-periodic microstructures, especially to derive macroscopic equations for problems posed in domains with perforations distributed non-periodically. Using the methods of locally periodic two-scale convergence (l-t-s) on oscillating surfaces and the locally periodic (l-p) boundary unfolding operator, we are able to analyze differential equations defined on boundaries of non-periodic microstructures and consider non-homogeneous Neumann conditions on the boundaries of perforations, distributed non-periodically.

In this paper, we extend previous work on the mathematical modeling of oxygen transport in biological tissues (Matzavinos et al., 2009). Specifically, we include in the modeling process the arterial and venous microstructure within the tissue by means of homogenization techniques. We focus on the two-layer tissue architecture investigated in (Matzavinos et al., 2009) in the context of abdominal tissue flaps that are commonly used for reconstructive surgery. We apply two-scale convergence methods and unfolding operator techniques to homogenize the developed microscopic model, which involves different unit-cell geometries in the two distinct tissue layers (skin layer and fat tissue) to account for different arterial branching patterns.

Gene regulatory networks, i.e. DNA segments in a cell which interact with each other indirectly through their RNA and protein products, lie at the heart of many important intracellular signal transduction processes. In this paper we analyse a mathematical model of a canonical gene regulatory network consisting of a single negative feedback loop between a protein and its mRNA (e.g. the Hes1 transcription factor system). The model consists of two partial differential equations describing the spatio-temporal interactions between the protein and its mRNA in a 1-dimensional domain. Such intracellular negative feedback systems are known to exhibit oscillatory behaviour and this is the case for our model, shown initially via computational simulations. In order to investigate this behaviour more deeply, we next solve our system using Green's functions and then undertake a linearized stability analysis of the steady states of the model. Our results show that the diffusion coefficient of the protein/mRNA acts as a bifurcation parameter and gives rise to a Hopf bifurcation. This shows that the spatial movement of the mRNA and protein molecules alone is sufficient to cause the oscillations. This has implications for transcription factors such as p53, NF-$\kappa$B and heat shock proteins which are involved in regulating important cellular processes such as inflammation, meiosis, apoptosis and the heat shock response, and are linked to diseases such as arthritis and cancer.

Regulation of zinc uptake in roots of Arabidopsis thaliana has recently been modeled by a system of ordinary differential equations based on the uptake of zinc, expression of a transporter protein and the interaction between an activator and inhibitor. For certain parameter choices the steady state of this model becomes unstable upon variation in the external zinc concentration. Numerical results show periodic orbits emerging between two critical values of the external zinc concentration. Here we show the existence of a global Hopf bifurcation with a continuous family of stable periodic orbits between two Hopf bifurcation points. The stability of the orbits in a neighborhood of the bifurcation points is analyzed by deriving the normal form, while the stability of the orbits in the global continuation is shown by calculation of the Floquet multipliers. From a biological point of view, stable periodic orbits lead to potentially toxic zinc peaks in plant cells. Buffering is believed to be an efficient way to deal with strong transient variations in zinc supply. We extend the model by a buffer reaction and analyze the stability of the steady state in dependence of the properties of this reaction. We find that a large enough equilibrium constant of the buffering reaction stabilizes the steady state and prevents the development of oscillations. Hence, our results suggest that buffering has a key role in the dynamics of zinc homeostasis in plant cells., Comment: 22 pages, 5 figures, uses svjour3.cls

In this paper, we focus on the Keller-Segel chemotaxis system in a random heterogeneous domain. We assume that the corresponding diffusion and chemotaxis coefficients are given by stationary ergodic random fields and apply stochastic two-scale convergence methods to derive the homogenized macroscopic equations. In establishing our results, we also derive a priori estimates for the Keller-Segel system that rely only on the boundedness of the coefficients; in particular, no differentiability assumption on the diffusion and chemotaxis coefficients for the chemotactic species is required. Finally, we prove the convergence of a periodization procedure for approximating the homogenized macroscopic coefficients.

Water flow in plant tissues takes place in two different physical domains separated by semipermeable membranes: cell insides and cell walls. The assembly of all cell insides and cell walls are termed symplast and apoplast, respectively. Water transport is pressure driven in both, where osmosis plays an essential role in membrane crossing. In this paper, a microscopic model of water flow and transport of an osmotically active solute in a plant tissue is considered. The model is posed on the scale of a single cell and the tissue is assumed to be composed of periodically distributed cells. The flow in the symplast can be regarded as a viscous Stokes flow, while Darcy's law applies in the porous apoplast. Transmission conditions at the interface (semipermeable membrane) are obtained by balancing the mass fluxes through the interface and by describing the protein mediated transport as a surface reaction. Applying homogenization techniques, macroscopic equations for water and solute transport in a plant tissue are derived. The macroscopic problem is given by a Darcy law with a force term proportional to the difference in concentrations of the osmotically active solute in the symplast and apoplast; i.e. the flow is also driven by the local concentration difference and its direction can be different than the one prescribed by the pressure gradient., Comment: 31 pages

In this paper we study pattern formation arising in a system of a single reaction-diffusion equation coupled with subsystem of ordinary differential equations, describing spatially-distributed growth of clonal populations of precancerous cells, whose proliferation is controled by growth factors diffusing in the extracellular medium and binding to the cell surface. We extend the results on the existence of nonhomogenous stationary solutions obtained in the previous work of one of the authors to a general Hill-type production function and full parameter set. Using spectral analysis and perturbation theory we derive conditions for the linearized stability of such spatial patterns., Comment: published in CPAA

We use the periodic unfolding technique to derive corrector estimates for a reaction-diffusion system describing concrete corrosion penetration in the sewer pipes. The system, defined in a periodically-perforated domain, is semi-linear, partially dissipative, and coupled via a non-linear ordinary differential equation posed on the solid-water interface at the pore level. After discussing the solvability of the pore scale model, we apply the periodic unfolding techniques (adapted to treat the presence of perforations) not only to get upscaled model equations, but also to prepare a proper framework for getting a convergence rate (corrector estimates) of the averaging procedure., Comment: 23 pages, one figure

The introduced notion of locally-periodic two-scale convergence allows to average a wider range of microstructures, compared to the periodic one. The compactness theorem for the locally-periodic two-scale convergence and the characterisation of the limit for a sequence bounded in $H^1(\Omega)$ are proven. The underlying analysis comprises the approximation of functions, which periodicity with respect to the fast variable depends on the slow variable, by locally-periodic functions, periodic in subdomains smaller than the considered domain, but larger than the size of microscopic structures. The developed theory is applied to derive macroscopic equations for a linear elasticity problem defined in domains with plywood structures., Comment: 22 pages, 4 figures