<p>I detta tvärvetenskapliga arbete studeras från den matematiska sidan tre klassiska populationsmodeller: Malthus tillväxtmodell, Verhulsts logistiska modell och Lotka-Volterras jägarebytesmodell. De klassiska modellerna jämförs med stokastiska. De stokastiska modeller som studeras är födelsedödsprocesser och deras diffusionsapproximation. Jämförelse görs med medelvärdesbildade simuleringar.</p><p>Det krävs många simuleringar för att kunna genomföra jämförelserna. Dessa simuleringar måste utföras i datormiljö och det är här den datalogiska aspekten av arbetet kommer in. Modellerna och deras resultathantering har implementerats i både MatLab och i C, för att kunna möjliggöra en undersökning om skillnaderna i tidsåtgången mellan de båda språken, under genomförandet av ovan nämnda jämförelser. Försök till tidsoptimering utförs och även användarvänligheten under implementeringen av de matematiska problemen i de båda språken behandlas.</p><p>Följande matematiska slutsatser har dragits, att de medelvärdesbildade lösningarna inte alltid sammanfaller med de klassiska modellerna när de simuleras på stora tidsintervall. I den logistiska modellen samt i Lotka-Volterras modell dör förr eller senare de stokastiska simuleringarna ut när tiden går mot oändligheten, medan deras deterministiska representation lever vidare. I den exponentiella modellen sammanfaller medelvärdet av de stokastiska simuleringarna med den deterministiska lösningen, dock blir spridningen stor för de stokastiska simuleringarna när de utförs på stora tidsintervall.</p><p>Datalogiska slutsatser som har dragits är att när det kommer till att implementera få modeller, samt resultatbearbetning av dessa, som ska användas upprepade gånger, är C det bäst lämpade språket då det visat sig vara betydligt snabbare under exekvering än vad MatLab är. Dock måste hänsyn tas till alla de svårigheter som implementeringen i C drar med sig. Dessa svårigheter kan till stor del undvikas om implementeringen istället sker i MatLab, då det därmed finns tillgång till en uppsjö av väl lämpade funktioner och färdiga matematiska lösningar.</p> / <p>In this interdisciplinary study, three classic population models will be studied from a mathematical view: Malthus’ growth, Verhulst’s logistic model and Lotka-Volterra’s model for hunter and prey. The classic models are being compared to the stochastic ones. The stochastic models studied are the birthdeath processes and their diffusion approximation. Comparisons are made by averaging simulations.</p><p>It requires numerous simulations to carry out the comparisons. The simulations must be carried out on a computer and this is where the computer science emerges to the project. The models, along with the handling of the results, have been implemented in both Mat- Lab and in C in order to allow a comparison between the two languages whilst executing the above mentioned study. Attempts to time optimization and an evaluation concerning the user-friendliness regarding the implementation of mathematical problems will be performed.</p><p>Mathematic conclusions, which have been drawn, are that the averaging solutions do not always coincide with the traditional models when they are being simulated over large time. In the logistic model and in Lotka-Volterra’s model the stochastic simulations will sooner or later die when the time is moving towards infinity, whilst their deterministic representation keeps on living. In the exponential model, the mean values of the stochastic simulations and of the deterministic solution coincide. There is, however, a large spread for the stochastic simulations when they are carried out over a large time.</p><p>Computer scientific conclusions drawn from the study includes that when it comes to implementing a few models, along with the handling of the results, to be used repeatedly, C is the most appropriate language as it proved to be significantly faster during execution. However, all of the difficulties during the implementation of mathematical problems in C must be kept in mind. These difficulties can be avoided if the implementation instead takes place in MatLab, where a numerous of mathematical functions and solutions will be available.</p>

<p>I detta tvärvetenskapliga arbete studeras från den matematiska sidan tre klassiska populationsmodeller: Malthus tillväxtmodell, Verhulsts logistiska modell och Lotka-Volterras jägarebytesmodell. De klassiska modellerna jämförs med stokastiska. De stokastiska modeller som studeras är födelsedödsprocesser och deras diffusionsapproximation. Jämförelse görs med medelvärdesbildade simuleringar.</p><p>Det krävs många simuleringar för att kunna genomföra jämförelserna. Dessa simuleringar måste utföras i datormiljö och det är här den datalogiska aspekten av arbetet kommer in. Modellerna och deras resultathantering har implementerats i både MatLab och i C, för att kunna möjliggöra en undersökning om skillnaderna i tidsåtgången mellan de båda språken, under genomförandet av ovan nämnda jämförelser. Försök till tidsoptimering utförs och även användarvänligheten under implementeringen av de matematiska problemen i de båda språken behandlas.</p><p>Följande matematiska slutsatser har dragits, att de medelvärdesbildade lösningarna inte alltid sammanfaller med de klassiska modellerna när de simuleras på stora tidsintervall. I den logistiska modellen samt i Lotka-Volterras modell dör förr eller senare de stokastiska simuleringarna ut när tiden går mot oändligheten, medan deras deterministiska representation lever vidare. I den exponentiella modellen sammanfaller medelvärdet av de stokastiska simuleringarna med den deterministiska lösningen, dock blir spridningen stor för de stokastiska simuleringarna när de utförs på stora tidsintervall.</p><p>Datalogiska slutsatser som har dragits är att när det kommer till att implementera få modeller, samt resultatbearbetning av dessa, som ska användas upprepade gånger, är C det bäst lämpade språket då det visat sig vara betydligt snabbare under exekvering än vad MatLab är. Dock måste hänsyn tas till alla de svårigheter som implementeringen i C drar med sig. Dessa svårigheter kan till stor del undvikas om implementeringen istället sker i MatLab, då det därmed finns tillgång till en uppsjö av väl lämpade funktioner och färdiga matematiska lösningar.</p> / <p>In this interdisciplinary study, three classic population models will be studied from a mathematical view: Malthus’ growth, Verhulst’s logistic model and Lotka-Volterra’s model for hunter and prey. The classic models are being compared to the stochastic ones. The stochastic models studied are the birthdeath processes and their diffusion approximation. Comparisons are made by averaging simulations.</p><p>It requires numerous simulations to carry out the comparisons. The simulations must be carried out on a computer and this is where the computer science emerges to the project. The models, along with the handling of the results, have been implemented in both MatLab and in C in order to allow a comparison between the two languages whilst executing the above mentioned study. Attempts to time optimization and an evaluation concerning the user-friendliness regarding the implementation of mathematical problems will be performed.</p><p>Mathematic conclusions, which have been drawn, are that the averaging solutions do not always coincide with the traditional models when they are being simulated over large time. In the logistic model and in Lotka-Volterra’s model the stochastic simulations will sooner or later die when the time is moving towards infinity, whilst their deterministic representation keeps on living. In the exponential model, the mean values of the stochastic simulations and of the deterministic solution coincide. There is, however, a large spread for the stochastic simulations when they are carried out over a large time.</p><p>Computer scientific conclusions drawn from the study includes that when it comes to implementing a few models, along with the handling of the results, to be used repeatedly, C is the most appropriate language as it proved to be significantly faster during execution. However, all of the difficulties during the implementation of mathematical problems in C must be kept in mind. These difficulties can be avoided if the implementation instead takes place in MatLab, where a numerous of mathematical functions and solutions will be available.</p>

I detta tvärvetenskapliga arbete studeras från den matematiska sidan tre klassiska populationsmodeller: Malthus tillväxtmodell, Verhulsts logistiska modell och Lotka-Volterras jägarebytesmodell. De klassiska modellerna jämförs med stokastiska. De stokastiska modeller som studeras är födelsedödsprocesser och deras diffusionsapproximation. Jämförelse görs med medelvärdesbildade simuleringar. Det krävs många simuleringar för att kunna genomföra jämförelserna. Dessa simuleringar måste utföras i datormiljö och det är här den datalogiska aspekten av arbetet kommer in. Modellerna och deras resultathantering har implementerats i både MatLab och i C, för att kunna möjliggöra en undersökning om skillnaderna i tidsåtgången mellan de båda språken, under genomförandet av ovan nämnda jämförelser. Försök till tidsoptimering utförs och även användarvänligheten under implementeringen av de matematiska problemen i de båda språken behandlas. Följande matematiska slutsatser har dragits, att de medelvärdesbildade lösningarna inte alltid sammanfaller med de klassiska modellerna när de simuleras på stora tidsintervall. I den logistiska modellen samt i Lotka-Volterras modell dör förr eller senare de stokastiska simuleringarna ut när tiden går mot oändligheten, medan deras deterministiska representation lever vidare. I den exponentiella modellen sammanfaller medelvärdet av de stokastiska simuleringarna med den deterministiska lösningen, dock blir spridningen stor för de stokastiska simuleringarna när de utförs på stora tidsintervall. Datalogiska slutsatser som har dragits är att när det kommer till att implementera få modeller, samt resultatbearbetning av dessa, som ska användas upprepade gånger, är C det bäst lämpade språket då det visat sig vara betydligt snabbare under exekvering än vad MatLab är. Dock måste hänsyn tas till alla de svårigheter som implementeringen i C drar med sig. Dessa svårigheter kan till stor del undvikas om implementeringen istället sker i MatLab, då det därmed finns tillgång till en uppsjö av väl lämpade funktioner och färdiga matematiska lösningar. / In this interdisciplinary study, three classic population models will be studied from a mathematical view: Malthus’ growth, Verhulst’s logistic model and Lotka-Volterra’s model for hunter and prey. The classic models are being compared to the stochastic ones. The stochastic models studied are the birthdeath processes and their diffusion approximation. Comparisons are made by averaging simulations. It requires numerous simulations to carry out the comparisons. The simulations must be carried out on a computer and this is where the computer science emerges to the project. The models, along with the handling of the results, have been implemented in both Mat- Lab and in C in order to allow a comparison between the two languages whilst executing the above mentioned study. Attempts to time optimization and an evaluation concerning the user-friendliness regarding the implementation of mathematical problems will be performed. Mathematic conclusions, which have been drawn, are that the averaging solutions do not always coincide with the traditional models when they are being simulated over large time. In the logistic model and in Lotka-Volterra’s model the stochastic simulations will sooner or later die when the time is moving towards infinity, whilst their deterministic representation keeps on living. In the exponential model, the mean values of the stochastic simulations and of the deterministic solution coincide. There is, however, a large spread for the stochastic simulations when they are carried out over a large time. Computer scientific conclusions drawn from the study includes that when it comes to implementing a few models, along with the handling of the results, to be used repeatedly, C is the most appropriate language as it proved to be significantly faster during execution. However, all of the difficulties during the implementation of mathematical problems in C must be kept in mind. These difficulties can be avoided if the implementation instead takes place in MatLab, where a numerous of mathematical functions and solutions will be available.

Nesta dissertação, tratamos do estudo das bifurcações de um modelo bi-dimensional de presa-predador, que estende e aperfeiçoa o sistema de Lotka-Volterra. Tal modelo apresenta cinco parâmetros e uma função resposta não monotônica do tipo Holling IV: $$ \\left\\{\\begin \\dot=x(1-\\lambda x-\\frac{\\alpha x^2+\\beta x +1})\\\\ \\dot=y(-\\delta-\\mu y+\\frac{\\alpha x^2+\\beta x +1}) \\end ight. $$ Estudamos as bifurcações do tipo sela-nó, Hopf, transcrítica, Bogdanov-Takens e Bogdanov-Takens degenerada. O método dos centros organizadores é usado para estudar o comportamento qualitativo do diagrama de bifurcação. / In this work are studied the bifurcations of a bi-dimensional predator-prey model, which extends and improves the Volterra-Lotka system. This model has five parameters and a non-monotonic response function of Holling IV type: $$ \\left\\{\\begin \\dot=x(1-\\lambda x-\\frac{\\alpha x^2+\\beta x +1})\\\\ \\dot=y(-\\delta-\\mu y+\\frac{\\alpha x^2+\\beta x +1}) \\end ight. $$ They studied the sadle-node, Hopf, transcritic, Bogdanov-Takens and degenerate Bogdanov-Takens bifurcations. The method of organising centers is used to study the qualitative behavior of the bifurcation diagram.

Traditionally, differential-equation models for population dynamics have considered organisms as "fixed" entities in terms of their behaviour and characteristics. However, there have been many observations of adaptivity in organisms, both at the level of behaviour and as an evolutionary change of traits, in response to the environmental conditions. Taking such adaptiveness into account alters the qualitative dynamics of traditional models and is an important factor to be included, for example, when developing reliable model predictions under changing environmental conditions. In this thesis, we consider piecewise-smooth and smooth dynamical systems to represent adaptive change in a 1 predator-2 prey system. First, we derive a novel piecewise-smooth dynamical system for a predator switching between its preferred and alternative prey type in response to prey abundance. We consider a linear ecological trade-off and discover a novel bifurcation as we change the slope of the trade-off. Second, we reformulate the piecewise-smooth system as two novel 1 predator-2 prey smooth dynamical systems. As opposed to the piecewise-smooth system that includes a discontinuity in the vector fields and assumes that a predator switches its feeding strategy instantaneously, we relax this assumption in these systems and consider continuous change in a predator trait. We use plankton as our reference organism because they serve as an important model system. We compare the model simulations with data from Lake Constance on the German-Swiss-Austrian border and suggest possible mechanistic explanations for cycles in plankton concentrations in spring.

This work is devoted to investigating the algebraic and analytic integrability of first order polynomial partial differential equations via an understanding of the well-developed area of local and global integrability of polynomial vector fields. In the view of characteristics method, the search of first integrals of the first order partial differential equations P(x,y,z)∂z(x,y) ∂x +Q(x,y,z)∂z(x,y) ∂y = R(x,y,z), (1) is equivalent to the search of first integrals of the system of the ordinary differential equations dx/dt= P(x,y,z), dy/dt= Q(x,y,z), dz/dt= R(x,y,z). (2) The trajectories of (2) will be found by representing these trajectories as the intersection of level surfaces of first integrals of (1). We would like to investigate the integrability of the partial differential equation (1) around a singularity. This is a case where understanding of ordinary differential equations will help understanding of partial differential equations. Clearly, first integrals of the partial differential equation (1), are first integrals of the ordinary differential equations (2). So, if (2) has two first integrals φ1(x,y,z) =C1and φ2(x,y,z) =C2, where C1and C2 are constants, then the general solution of (1) is F(φ1,φ2) = 0, where F is an arbitrary function of φ1and φ2. We choose for our investigation a system with quadratic nonlinearities and such that the axes planes are invariant for the characteristics: this gives three dimensional Lotka– Volterra systems x' =dx/dt= P = x(λ +ax+by+cz), y' =dy/dt= Q = y(µ +dx+ey+ fz), z' =dz/dt= R = z(ν +gx+hy+kz), where λ,µ,ν 6= 0. v Several problems have been investigated in this work such as the study of local integrability and linearizability of three dimensional Lotka–Volterra equations with (λ:µ:ν)–resonance. More precisely, we give a complete set of necessary and sufficient conditions for both integrability and linearizability for three dimensional Lotka-Volterra systems for (1:−1:1), (2:−1:1) and (1:−2:1)–resonance. To prove their sufficiency, we mainly use the method of Darboux with the existence of inverse Jacobi multipliers, and the linearizability of a node in two variables with power-series arguments in the third variable. Also, more general three dimensional system have been investigated and necessary and sufficient conditions are obtained. In another approach, we also consider the applicability of an entirely different method which based on the monodromy method to prove the sufficiency of integrability of these systems. These investigations, in fact, mean that we generalized the classical centre-focus problem in two dimensional vector fields to three dimensional vector fields. In three dimensions, the possible mechanisms underling integrability are more difficult and computationally much harder. We also give a generalization of Singer’s theorem about the existence of Liouvillian first integrals in codimension 1 foliations in Cnas well as to three dimensional vector fields. Finally, we characterize the centres of the quasi-homogeneous planar polynomial differential systems of degree three. We show that at most one limit cycle can bifurcate from the periodic orbits of a centre of a cubic homogeneous polynomial system using the averaging theory of first order.

Работа содержит исследования трех моделей живых систем с дискретным временем. В первой главе рассматривается одномерная модель нейронной активности, задаваемая кусочно-гладким отображением. Показывается, что в случае одномерного отображения наличие случайного возмущения приводит к появлению всплесков (спайкингу). Исследуются два механизма генерации спайков, вызванных добавлением случайного возмущения в один из параметров. Иллюстрируется, что сосуществование двух аттракторов является не единственной причиной возникновения спайкинга. Для прогнозирования уровня интенсивности шума, необходимого для генерации спайков, применяется метод доверительных областей, который основан на функции стохастической чувствительности. Также находятся основные характеристики межспайковых интервалов в зависимости от интенсивности шума. Вторая глава работы посвящена применению метода функции стохастической чувствительности к аттракторам кусочно-гладкого одномерного отображения, описывающего динамику численности популяции. Первым этапом исследования является параметрический анализ возможных режимов детерминированной модели: определение зон существования устойчивых равновесий и хаотических аттракторов. Для определения параметрических границ хаотического аттрактора применяется теория критических точек. В случае, когда на систему оказывает влияние случайное воздействие, на основе техники функции стохастической чувствительности дается описание разброса случайных состояний вокруг равновесия и хаотического аттрактора. Проводится сравнительный анализ влияния параметрического и аддитивного шума на аттракторы системы. С помощью техники доверительных интервалов изучаются вероятностные механизмы вымирания популяции под действием шума. Анализируются изменения параметрических границ существования популяции под действием случайного возмущения. В третьей главе проводится анализ возможных динамических режимов детерминированной и стохастической модели Лотки-Вольтерры. В зависимости от двух параметров системы строится карта режимов. Изучаются параметрические зоны существования устойчивых равновесий, циклов, замкнутых инвариантных кривых, а также хаотических аттракторов. Описываются бифуркации удвоения периода, Неймарка--Саккера и кризиса. Демонстрируется сложная форма бассейнов притяжения. Помимо детерминированной системы подробно изучается стохастическая, описывающая влияние внешнего случайного воздействия. В случае хаоса дан алгоритм нахождения критических линий, описывающих границу хаотического аттрактора. Опираясь на найденную чувствительность аттракторов, строятся доверительные полосы и эллипсы, позволяющие описать разброс случайных состояний вокруг детерминированного аттрактора. / The work contains study of three models of biological systems with discrete time. In the first chapter a one-dimensional model of neural activity defined by a piecewise-smooth map is considered. It is shown that in the case of a one-dimensional model, the presence of a random disturbance leads to a spike generation. Two mechanisms of spike generation caused by the presence of a random disturbance in one of the parameters are investigated. It is illustrated that the coexistence of two attractors is not the only reason of spiking. To predict the level of noise intensity needed to generate spikes, the confidence-domain method is used, which is based on the stochastic sensitivity function. The main characteristics of interspike intervals depending on the intensity of the noise are also described. The second chapter is devoted to the application of the method of the stochastic sensitivity function to attractors of a piecewise-smooth one-dimensional map, which describes the population dynamics. The first stage of the study is a parametric analysis of the possible regimes of the deterministic model: determining the zones of existence of stable equilibria and chaotic attractors. The theory of critical points is used to determine the parametric boundaries of a chaotic attractor. In the case where the system is affected by a random noise, based on the stochastic sensitivity function, a description of the spread of random states around equilibrium and a chaotic attractor is given. A comparative analysis of the influence of parametric and additive noise on the attractors is carried out. Using the technique of confidence intervals, the probabilistic mechanisms of extinction of a population under the influence of noise are studied. Changes in the parametric boundaries of the existence of population under the influence of random disturbance are analyzed. In the third chapter the possible dynamic modes of the Lotka-Volterra model in determi\-nistic and stochastic cases are analyzed. Depending on the two parameters of the system, bifurcation diagram is constructed. Parametric zones of the existence of stable equilibria, cycles, closed invariant curves, and also chaotic attractors are studied. The bifurcations of the period doubling, Neimark--Sacker and the crisis are described. The complex shape of the basins of attraction is demonstrated. In addition to the deterministic system, the stochastic system is studied in detail, which describes the influence of external random disturbance. In the case of chaos, an algorithm for finding critical lines describing the boundary of a chaotic attractor is given. Based on the stochastic sensitivity function, confidence bands and ellipses are constructed to describe the spread of random states around a deterministic attractor.

Biological invasions provide a unique opportunity to study the mechanisms that regulate community composition and ecosystem function. Invasive species that are also ecosystem engineers can substantially alter physical features in an environment, and this can lead to cascading effects on the biological community. Aquatic-terrestrial interface ecosystems are excellent systems to study the interactions among invasive ecosystem engineers, physical features, and biological communities, because interactions among vegetation, sediment, and fluids within biophysical feedbacks create and modify distinct physical features. Further, these systems provide important ecosystem services including coastal protection afforded by their natural features. In this dissertation, I investigate the interactions and feedbacks among sand-binding beach grass species (a native, Elymus mollis (Trin.), and two non-natives, Ammophila arenaria (L.) Link and A. breviligulata Fernald), sediment supply, and dune shape along the U.S. Pacific Northwest coast. Dunes dominated by A. arenaria tend to be taller and narrower compared to the shorter, wider dunes dominated by A. breviligulata. These patterns suggest an ecological control on dune shape, and thus, coastal vulnerability to overtopping waves. I investigate the causes and consequences of these patterns with experiments, field observations, and modeling. Specifically, I investigate the relative roles of vegetation and sediment supply in shaping coastal dunes over inter-annual and multi-decadal time scales (Chapter 2), characterize a biophysical feedback between beach grass species growth habit and sediment supply (Chapter 3), uncover the mechanisms leading to beach grass coexistence and whether A. breviligulata can invade and dominate new sections of coastline (Chapter 4), and examine the non-target effects resulting from management actions that remove Ammophila for the recovery of the threatened Western Snowy plover (Charadrius alexandrinus nivosus) (Chapter 5). I found that vegetation and sediment supply play important roles in dune shape changes across inter-annual and multi-decadal time scales (Chapter 2). I determined that a biophysical feedback between the beach grass growth habits and sediment supply results in species-specific differences in sand capture ability, and thus, is a likely explanation for differences in dune shape (Chapter 3). I found that all three beach grass species can coexist across different sediment deposition rates, and that this coexistence is largely mediated by positive direct and indirect species interactions. I further determined that A. breviligulata is capable of invading and dominating the beach grass community in regions where it is currently absent (Chapter 4). Combined, these findings indicate that A. breviligulata is an inferior dune building species as compared to A. arenaria, and suggest that in combination with sediment supply gradients, these species differences ultimately lead to differences in dune shape. Potential further invasions of A. breviligulata into southern regions of the Pacific Northwest may diminish the coastal protection ability of dunes currently dominated by A. arenaria, but this effect could be moderated by the predicted near co-dominance of A. arenaria in these lower sediment supply conditions. Finally, I found that the techniques used to remove Ammophila for plover recovery have unintended consequences for the native and endemic dune plant communities, and disrupt the natural disturbance regime of shifting sand. A whole-ecosystem restoration focus would be an improvement over the target-species approach, as it would promote the return of the natural disturbance regime, which in turn, would help recover the native biological community. The findings from this dissertation research provide a robust knowledge base that can guide further investigations of biological and physical changes to the coastal dunes, can help improve the management of dune ecosystem services and the restoration of native communities, and can help anticipate the impacts of future beach grass invasions and climate change induced changes to the coast. / Graduation date: 2012 / Access restricted to the OSU Community at author's request from Sept. 22, 2011 - March 22, 2012