Rapid prototyping tools exist in many fields of science and engineering, but are rare in biology especially not general tools that can handle the diversity and complexity of the many spatial and temporal scales in nature. In this thesis a general use, cell-based, middle-out biology emulation programming framework (outlining a programming paradigm) is presented, that enables biologists to emulate and use virtual biological systems of previously unimaginable complexity and potentially get results accurate enough to be used in research and ultimately, in clinical practice, such as diagnosis or operations. With this technology, virtual organisms can be created that are viable, fit and can be optimised for any task that arises. The tool, realised with a programming framework created for the C++ language is detailed and demonstrated through several examples of increasing complexity, namely several example organisms and a cancer emulation, showing both viable virtual organisms and usable experimental results.

Embryonic development depends on the precise coordination of cell fate specification, patterning and morphogenesis. Although great strides have been made in the molecular understanding of each of these processes, how their interplay governs the formation of complex tissues remains poorly understood. New techniques for experimental manipulation and image quantification enable the study of development in unprecedented detail, resulting in new hypotheses on the interactions between known components. By expressing these hypotheses in terms of rules and equations, computational modeling and simulation allows one to test their consistency against experimental data. However, new computational methods are required to represent and integrate the network of interactions between gene regulation, signaling and biomechanics that extend over the molecular, cellular and tissue scales. In this thesis, I present a framework that facilitates computational modeling of multiscale multicellular systems and apply it to investigate pancreatic development and the formation of vascular networks. This framework is based on the integration of discrete cell-based models with continuous models for intracellular regulation and intercellular signaling. Specifically, gene regulatory networks are represented by differential equations to analyze cell fate regulation; interactions and distributions of signaling molecules are modeled by reaction-diffusion systems to study pattern formation; and cell-cell interactions are represented in cell-based models to investigate morphogenetic processes. A cell-centered approach is adopted that facilitates the integration of processes across the scales and simultaneously constrains model complexity. The computational methods that are required for this modeling framework have been implemented in the software platform Morpheus. This modeling and simulation environment enables the development, execution and analysis of multi-scale models of multicellular systems. These models are represented in a new domain-specific markup language that separates the biological model from the computational methods and facilitates model storage and exchange. Together with a user-friendly graphical interface, Morpheus enables computational modeling of complex developmental processes without programming and thereby widens its accessibility for biologists. To demonstrate the applicability of the framework to problems in developmental biology, two case studies are presented that address different aspects of the interplay between cell fate specification, patterning and morphogenesis. In the first, I focus on the interplay between cell fate stability and intercellular signaling. Specifically, two studies are presented that investigate how mechanisms of cell-cell communication affect cell fate regulation and spatial patterning in the pancreatic epithelium. Using bifurcation analysis and simulations of spatially coupled differential equations, it is shown that intercellular communication results in a multistability of gene expression states that can explain the scattered spatial distribution and low cell type ratio of nascent islet cells. Moreover, model analysis shows that disruption of intercellular communication induces a transition between gene expression states that can explain observations of in vitro transdifferentiation from adult acinar cells into new islet cells. These results emphasize the role of the multicellular context in cell fate regulation during development and may be used to optimize protocols for cellular reprogramming. The second case study focuses on the feedback between patterning and morphogenesis in the context of the formation of vascular networks. Integrating a cell-based model of endothelial chemotaxis with a reaction-diffusion model representing signaling molecules and extracellular matrix, it is shown that vascular network patterns with realistic morphometry can arise when signaling factors are retained by cell-modified matrix molecules. Through the validation of this model using in vitro assays, quantitative estimates are obtained for kinetic parameters that, when used in quantitative model simulations, confirm the formation of vascular networks under measured biophysical conditions. These results demonstrate the key role of the extracellular matrix in providing spatial guidance cues, a fact that may be exploited to enhance vascularization of engineered tissues. Together, the modeling framework, software platform and case studies presented in this thesis demonstrate how cell-centered computational modeling of multi-scale and multicellular systems provide powerful tools to help disentangle the complex interplay between cell fate specification, patterning and morphogenesis during embryonic development.

Les agrégats multi-échelles (sulfure de zinc: ZnS, ...) ont des propriétés physiques et physicochimiques présentant un grand intérêt industriel (industrie pharmaceutique, matériau, environnement...). L'objectif général de cette thèse est de caractériser ces agrégats multi-échelles par une méthode optique d'une part, et par une étude géométrique d'autre part. Le but est de trouver des relations simples entre propriétés optiques et géométriques. Les résultats présentés sont valables pour tout matériau à haut indice de réfraction.Ce travail comporte une première étape de modélisation en étudiant tout d'abord des modèles simples d'agrégats multi-échelles (agencement ordonné de particules élémentaires sphériques). Différents agrégats sont alors géométriquement modélisés suivant des paramètres de construction prédéfinis (volume global des agrégats, rayon des particules primaires...). Une analyse préliminaire optique puis une analyse géométrique sont alors effectuées afin de déterminer les paramètres respectifs à étudier: la section efficace moyenne de diffusion (...), calculée au moyen d'une extension de la théorie de Mie (GMM pour Generalized Multiparticle Mie) d'Y-L Xu, et des caractéristiques géométriques spécifiques (volume de matière, compacité...).Une seconde étape consiste à étudier les variations de la valeur de ... en fonction de celles des paramètres géométriques des agrégats. Des relations caractérisant ces variations sont établies. Enfin, une expression approchées pour la valeur de ... est recherchée sous la forme d'une loi de puissance, reposant sur des méthodes optiques approchées existantes et sur la sélection de caractéristiques géométriques pertinentes. Le modèle mis en place est robuste et présente un compromis entre précision et complexité. Le travail réalisé ouvre diverses perspectives comme la confrontation du modèle mathématique théorique obtenu, avec l'expérience. / Multiscale aggregates (zinc sulfide: ZnS, ...) havephysical and physicochemical properties of great industrial interest (e.g. pharmaceutical industriy, material, environment). The overall objective of this thesis is to characterize these multiscale aggregates by an optical method on the one hand, and a geometric study of the other. The goal is to find simple relations between optical and geometrical properties. The results presented are valid for any material with high refractive index.The first step of this work is to build in a simple way the multiscale aggregate (like ordered arrangement of spherical elementary particles). Different aggregates are then geometrically modeled thanks to predefined design parameters (e.g. particles radius, aggregates overall volume). A preleminary analysis of optical and geometrical analysis is then performed to determine the relevant parameters: the average scattering cross section (...), calculed using a generalization of the Mie theory (GMM for Generalized Multiparticle Mie) developped by Y.L. Xu, and specific geometric values features. An analysis of the behavior of the optical parameter and the geometrical characteristic (e.g. compactness, volume of matter...), with respect to the construction parameters of the aggregates, is then performed.A second step is to study variations of ... value versus the variation of the geometrical parameter of aggregates. Mathematical relations characterizing these variations are established. Finally, a study on the approximation of the ... value is carried out. A power law is proposed which is basedon previous optical approximate methods and on the selection of geometrical characteristics of the aggregates. The optical properties and geometrical characteristics of the clusters are fairly extensive and contrasted to give a general groundwork to this study. The work done opens various perspectives such as the confrontation of the theorical mathematical model obtained, with experiment.

Dislocation channel-veins and persistent slip bands (PSBs) are characteristic dislocation configurations that are of interest to both industry and academia. However, existing mathematical models are not adequate to describe the mechanism of the transition between these two states. In this thesis, a series of models are proposed to give a quantitative description to such a transition. The full problem has been considered from two angles. Firstly, the general motion and instabilities of arbitrary curved dislocations have been studied both analytically and numerically. Then the law of motion and local expansions are used to track the shapes of screw segments moving through channels, which are believed to induce dislocation multiplication by cross-slip. The second approach has been to investigate the collective behavior of a large number of dislocations, both geometrically necessary and otherwise. The traditional method of multiple scales does not apply well to describe the pile-up of two arrays of dislocations of opposite signs on a pair of neighbouring glide planes in two dimensional space. Certain quantities have to be more accurately defined under the multiple-scale coordinates to capture the much more localised resultant stress caused by these dislocation pairs. Through detailed calculations, one-dimensional dipoles can be homogenised to obtain some insightful results both on a local scale where the dipole pattern is the key diagnostic and on a macroscopic scale on which density variations are of most interest. Equilibria of dislocation dipoles in a two-dimensional regular lattice have been also studied. Some natural transitions between different patterns can be found as a result of geometrical instabilities.

Les matériaux composites sont largement utilisés dans les structures aéronautiques. Les travaux présentés ici visent à mettre en place une méthode de calcul permettant de prédire les délaminages dans ces structures stratifiées soumises à des chargements dynamiques tout en assurant des coûts de calculs compatibles dans un contexte industriel. Une méthode de décomposition de domaine en dynamique est d’abord utilisée, afin de coupler des modélisations et des pas de temps de calcul différents. Une modélisation fine est utilisée uniquement dans les zones en cours de dégradation. Une représentation macroscopique du stratifié par des éléments de coque 3D est développée et employée dans le reste de la structure. Les coûts de calcul sont ainsi réduits tout en assurant une bonne précision des résultats. Pour éviter un remaillage avec l’avancée du délaminage, une approche multi-échelle en temps et en espace est ensuite proposée. Un maillage global de coque 3D de l'ensemble de la structure est défini. Des maillages utilisant une représentation fine du matériau sont utilisés localement. Des pas de temps adaptés sont utilisés dans les deux types de maillage. / The part of composite materials in aeronautic structures is increasing due to their specific properties and the mass reduction they enable. Accurate numerical simulations are thus needed in order to design these structures, particularly to verify if they could resist dynamic charges such as soft bodies impact. Nevertheless, using a refined model to represent phenomenon such as delamination leads to computing time and dofs number incompatible with an industrial context. The aim of the present work is to propose a multi-scale method in space and time to solve dynamic impact problems on laminate structures. A domain decomposition method for dynamic problems is first used to couple different kinds of models and time discretisations. A refined model for the laminate is used in the degradating areas only - elsewhere, a coarser representation using 3D-shell elements is used. This approach reduced the cost of the simulation giving accurate results. To avoid a remeshing due to delamination propagation within the structure, a multi-scale method is then proposed. A global 3D shell elements mesh for the whole structure is defined. Local meshes based on a refined representation of the laminates are used only if required. Coupling between global and local representation is done using velocity field.

L’objectif de ce travail de thèse est l’étude des chocs pyrotechnique à la source par une approche fréquentielle et de leur simulation à l’aide de la TVRC (Théorie Variationnelle des Rayons Complexes). Cette étude s’appuie largement sur un essai réalisé par le CNES dans le cadre du pôle chocs pyrotechniques en juin 2006. Afin de traiter le problème posé par la simulation de la propagation des chocs dans le démonstrateur, la TVRC a été étendue aux coques orthotropes de courbure quelconque. Une formulation variationnelle adaptée a été exhibée et la relation de dispersion a été établie. Les travaux effectués montrent que la prise en compte des moyennes et hautes fréquences, traditionnellement négligées, peut s’avérer essentielle pour la compréhension des phénomènes de propagation des ondes de flexion dans les structures complexes. / This memory is about the study of pyroshocks near the source by a frequency approach and their simulation by the mean of the VCTR (variational Theory of Complex Rays). This study is based on a test made by the CNES (French Space Agency) in the frame of the Pyroshock Pole in june 2006. In order to deal with the problem of shock propagation in the structure on which the test has been proceeded, the VTCR has been extended to orthotropic shells with arbitrary curvature. An adapted variational formulation and the dispersion relation have been established. Taking mid- and high-frequencies into account can be a key point in order to understand propagation phenomena of flexural waves in complex structures.

Les matériaux composites stratifiés sont de plus en plus utilisés dans l'aéronautique, mais ils peuvent être sujets à large délaminage si soumis à impact. La nécessité d'effectuer des simulations numériques pour prédire l’endommagement devient essentielle pour l’ingénieur. Dans ce contexte, l'utilisation d'une modélisation fine semble préférable. En revanche, le coût de calcul associé serait prohibitif pour larges structures. Le but de ce travail consiste à réduire ce coût de calcul, en couplant le modèle fin, restreint à la zone active de délaminage, avec un modèle grossier appliqué au reste de la structure. En raison du comportement transitoire des problèmes d'impact, l'adaptabilité dynamique des modèles pour suivre les phénomènes évolutifs représente un point crucial de la stratégie de couplage. Des méthodes avancées sont utilisées pour coupler différents modèles. Par exemple, la méthode de Décomposition de Domaines, appliquée à l'adaptabilité dynamique, doit être combinée avec une stratégie de remaillage, considérée comme intrusive pour la mise en œuvre d'un logiciel pour Analyse à Eléments Finis. Dans ce travail, les bases d'une approche faiblement intrusive, la méthode de Substitution, sont présentés dans le domaine de la dynamique explicite. Il s’agît d’une formulation globale-locale, conçue pour appliquer un modèle grossier sur tout le domaine pour obtenir une réponse globale: ce pré-calcul est ensuite corrigé itérativement par l'application du modèle raffiné appliqué seulement où nécessaire. La vérification de la méthode de Substitution en comparaison avec la méthode de Décomposition de Domaines est présentée. / Composite laminates are increasingly employed in aeronautics, but can be prone to extensive delamination when submitted to impact loads. The need of performing virtual testing to predict delamination becomes essential for engineering workflows, in which the use of a fine modeling scheme appears nowadays to be the preferred one. The associated computational cost would be prohibitively high for large structures. The goal of this work consists in reducing such computational cost coupling the fine model, restricted to the surroundings of the delamination process zone, with a coarse one applied to the rest of the structure. Due to the transient behavior of impact problems, the dynamic adaptivity of the models to follow evolutive phenomena represents a crucial feature for the coupling. Many methodologies are currently used to couple multiple models, such as non-overlapping Domain Decomposition method, that, applied to dynamic adaptivity, has to be combined with a re-meshing strategy, considered as intrusive implementation within a Finite Element Analysis software. In this work, the bases of a weakly-intrusive approach, called Substitution method, are presented in the field of explicit dynamics. The method is based on a global-local formulation and is designed so that it is possible to make use of the pre-fixed coarse model the meshes the whole structure to obtain a global response: this pre-computation is then iteratively corrected considering the application of the refined model only where required, in the picture of an adaptive strategy. The verification of the Substitution method in comparison with the Domain Decomposition method is presented.

Ce travail de thèse est dédié au développement d’un modèle numérique prédictif du comportement de tissu sec soumis à l’impact à haute vitesse. La capacité d’un modèle numérique à prédire au plus juste des performances balistiques d’un tissu est étroitement liée à la bonne représentation des énergies en jeu et de la dynamique de rupture des plis. Parmi les différentes stratégies adoptées pour modéliser un tissu, les modèles mésoscopiques sont les plus populaires du fait de leur capacité à représenter fidèlement l’évolution de l’impact combinée à un coût de calcul raisonnable ; les torons sont alors traités comme des milieux continus.Pour représenter un ensemble de fibres disjointes comme un milieu continu, un modèle constitutif approprié est nécessaire. Celui généralement adopté est capable de bien représenter les propriétés longitudinales d’un toron mais limitatif pour représenter le comportement de sa section droite. Récemment, différents travaux ont démontré l’importance de la mécanique liée à la section droite d’un toron dans la rupture des tissus. Elle joue un rôle majeur pour aboutir à un modèle numérique prédictif. L’objet de cette thèse est de développer une nouvelle loi constitutive capable de surpasser les limites du modèle linéaire élastique classique tout en maintenant une bonne représentation des propriétés longitudinales du toron et un coût de calcul acceptable.La première étape a été de comprendre les phénomènes physiques et de quantifier en particulier les effets liés à la section droite du toron sur les propriétés balistiques d’un tissu. Pour répondre à cette question, deux modèles numériques à l’échelle microscopique d’un toron de Kevlar soumis à l’impact transverse ont été développés. Les résultats obtenus ont montré que la mécanique de la section droite a des répercussions sur l’initialisation de la rupture du fil mettant en jeu des énergies significatives pendant la première phase d’un impact. La nécessité de prendre en compte les aspects physiques liés à la section droite d’un toron a été finalement confirmée.A partir des résultats précédents, un nouveau modèle constitutif de toron adapté à des applications dynamiques a été développé. Une formulation hyperélastique, précédemment utilisée pour des analyses statiques a été étendue au cas de l’impact et une nouvelle approche multi-échelle a été proposée pour la détermination des paramètres matériaux. La validation de la nouvelle loi a été faite en comparant les résultats obtenus à l’échelle mésoscopique (toron) avec ceux obtenus par les analyses à la micro-échelle (fibre). L’approche proposée est capable de reproduire l’évolution de la section droite du fil pendant l’impact en gardant la bonne représentation de ses propriétés longitudinales.Par la suite, le modèle de toron proposé a été implémenté au niveau du tissu. Les résultats ont confirmé les observations faites à l’échelle du toron. Le modèle de tissu composé par des torons hyperélastiques est capable de représenter correctement la dynamique d’impact , l’évolution des énergies en jeu et la rupture du tissu. La stabilité numérique du modèle a également pu être appréciée.Finalement, le modèle mésoscopique de toron proposé permet une bonne représentation du comportement dynamique et appréhension de la rupture. Il devient ainsi un outil pratique et efficace pour la prédiction des performances balistiques de tissus. / The current thesis work focused on the development of a predictive numerical model of dry fabrics under high velocity impact.A mature bibliography exists on the subject. The impact phenomenon can be essentially resumed as an energy transfer between the colliding object and the fabric layers. The correct prediction of the fabric ballistic performance by a numerical model is related to the correct representation of the fabric energy evolution and its failure dynamic. Different numerical strategies have been proposed to model a fabric under ballistic impact. Mesoscopic numerical models resulted to be the most popular since they provide a realistic representation of the phenomenon for a reasonable computational cost. This is possible thanks to the main assumption of treating yarns as continuous media.In order to represent a discrete fiber bundle as a continuum an appropriate constitutive behavior have to be formulated. The universally adopted constitutive law accurately describes yarns longitudinal properties but it is limited in the representation of their transverse mechanical behavior. Recent studies have demonstrated how this last point is intrinsically related to fabrics failure and multilayer textiles response, then its correct representation becomes a critical point for an accurate model. The goal of the current work has been to provide a new constitutive model which overcome the limitation of the classic linear elastic approach while keeping unaltered its advantages, i.e. low computational costs and accurate description of yarn longitudinal behavior.The first step of this dissertation was to quantify the yarn cross section effects over textile ballistic properties and the phenomena related to this aspect. In order to provide an answer, two microscopic numerical studies of a single Kevlar yarn transversely impacted have been presented. Results showed how yarn transverse mechanical behavior has a role in failure initialization, while its energetic contribution to the global energy balance is not negligible during the first phases of an impact. The importance of a correct representation of the yarn transverse behavior for a predictive fabric numerical model was then confirmed.Starting from the previous microscopic observations, a consistent yarn continuum model for impact applications has been proposed. An hyperelastic formulation previously developed for static applications has been extended to impact analyses and a novel multiscale approach for the determination of all the material parameters has been introduced. The validation of the hyperelastic approach has been performed comparing the results with those obtained at the microscale. Compared to the classical approach, the introduced constitutive law is actually able to reproduce the evolution of the yarn cross section during the impact while keeping a correct representation of the yarn longitudinal properties. Moreover, the proposed formulation provides new physical measurement to exploit the physic behind the impact and new possibilities in terms of failure modelisation.In the final part of the dissertation, the proposed yarn continuum model is introduced at the fabric level. Results confirmed the observation performed at the yarn level. The proposed hyperelastic approach is able to correctly represent the impact dynamic and fabric energies trends. Moreover, it provides more stability and a better representation of the fabric failure compared to linear elastic approach. The proposed hyperelastic constitutive law and the linear elastic one can be adopted for different portion of the same yarn without occurring into model instabilities and providing accurate results.The yarn mesoscopic model developed in the current work offers new possibilities in terms of failure modelisation and post processing tools. These could be used to develop more accurate fabric model and exploit the phenomena behind fabrics and yarns failure mechanic.

<div>Electronics have now become a part of our daily life and therefore the reliability of microelectronics cannot be overlooked. As the Moore's law era comes to an end, various new system-level innovations (e.g., 3D packaging, evolution of packaging material to molding compounds) with constant scaling of transistors have resulted in increasingly complicated integrated circuits (ICs) configurations. The reliability modeling of complex ICs is a nontrivial concern for a variety of reasons. For example, ever since 2004, self-heating effect (SHE) has become an important reliability concern for ICs. Currently, many groups have developed thermal predictive models for transistors, circuits, and systems. In order to describe SHE self-consistently, the modeling framework must account for correlated self-heating within the ICs. This multi-scales nature of the self-consistency problem is one of the difficult factors poses an important challenge to self-consistent modeling. In addition, coupling between different physical effects within IC further complicates the problem.</div><div><br></div><div>In this thesis, we discuss three challenges, and their solutions related to an IC's reliability issues. We (i) generalize the classical effective medium theory (EMT) to account for anisotropic, heterogeneous system; (ii) develop computationally efficient a physics-based thermal compact model for a packaged ICs to predict junction temperature in the transistor based on the EMT model, and image charge theory. Our thermal compact model bridges different length scales among the sources and rest of the system. Finally (iii) propose the modeling framework of electrical chip package interaction (CPI) due to charge transport within mold compounds by coupling moisture diffusion, electric distribution, and ions transport. The proposed modeling framework not only addresses the three major modeling challenges discussed earlier, but also provides deep and fundamental insights regarding the performance and reliability of modern ICs. </div>

O controle biológico de pragas agrícolas é um dos serviços ecossistêmicos mais valorizados, dada sua importância para a produção agrícola. Embora vários estudos tenham demonstrado que a abundância e riqueza de predadores aumentam com a quantidade de floresta nativa, os mecanismos subjacentes que modulam a relação entre a cobertura florestal e provisão de controle biológico em diferentes escalas espaciais são ainda pouco conhecidos. Neste trabalho, utilizamos experimentos de exclusão de vertebrados voadores em oito paisagens cafeeiras na Mata Atlântica, num gradiente de cobertura florestal, e quantificamos as consequências para a perda foliar e para a frutificação. A perda foliar mostrou uma relação negativa com a cobertura florestal em paisagens com 2 km de raio, indicando que a herbivoria é melhor controlada em paisagens com alta cobertura florestal, especialmente na presença de aves e morcegos. No entanto, no nível local, 300 m ao redor das plantas de café, a perda foliar e a frutificação responderam diferentemente à cobertura florestal. Em unidades com baixa cobertura florestal local, a exclusão de aves e morcegos aumentou a perda foliar e diminuiu a frutificação em uma média de 13%. Por outro lado, em unidades com alta cobertura florestal local, a exclusão de aves e morcegos não teve efeitos significativos nem na perda foliar, nem na frutificação. Concluímos que os efeitos da exclusão de aves e morcegos na perda foliar e frutificação são modulados por diferentes processos que ocorrem no nível local e da paisagem. Sugerimos que quando a cobertura florestal local é alta (geralmente perto de fragmentos florestais), as aves e os morcegos não se alimentam apenas de herbívoros, mas também de mesopredadores. No entanto, quando a cobertura florestal local é baixa (e.g. longe de fragmentos florestais), os mesopredadores não ocorrem e aves e morcegos passam a prover serviço de controle biológico de pragas, alimentando-se principalmente de herbívoros. Destacamos a importância de empregar uma análise multiescalar em sistemas onde espécies com diferentes capacidades de dispersão proveem um serviço ecossistêmico. / Biological control of agricultural pests is one of the most important ecosystem services given its key role for agricultural production. Although several studies have shown that the abundance and richness of predators increase with the amount of native forest in the landscape, the underlying mechanisms relating forest cover at different spatial scales with the provision of biological control are still poorly understood. We experimentally excluded flying vertebrates (birds and bats) in eight coffee landscapes in the Brazilian Atlantic Forest, encompassing a gradient of forest cover and quantified the consequences for coffee leaf loss and fruit set. Leaf loss showed a negative relation with forest cover at landscape level, in landscapes with 2 km radius, indicating that herbivory is better controlled in landscapes with high forest cover, especially in the presence of birds and bats. However, at local level, 300 m around coffee plants, leaf loss and fruit set responded to forest cover differently. In units with low local forest cover exclusion of birds and bats increased leaf loss and reduced fruit set by 13% in mean. However, in units with high local forest cover, exclusion of birds and bats had no significant effect neither on leaf loss nor in fruit set. We concluded that the effects of birds and bats exclusion on leaf loss and fruit set are modulated by different processes occurring at landscape and local levels. We hypothesized that when local forest cover is high (usually near remnant forest fragments) birds and bats are not only controlling herbivores but may also be reducing mesopredators, while when local forest cover is low (e.g. far from forest fragments), mesopredators do not occur and birds and bats start providing biological pest control by feeding mainly on herbivores. We highlight the importance of employing a multiscale analysis in systems where species with different dispersal abilities are providing an ecosystem service.