The spatio-temporal dynamics of neutral genetic diversity

International audienceThe notions of pulled and pushed solutions of reaction-dispersal equations introduced by Garnier et al. (2012) and Roques et al. (2012) are based on a decomposition of the solutions into several components. In the framework of population dynamics, this decomposition is related to the spatio-temporal evolution of the genetic structure of a population. The pulled solutions describe a rapid erosion of neutral genetic diversity, while the pushed solutions are associated with a maintenance of diversity. This paper is a survey of the most recent applications of these notions to several standard models of population dynamics, including reaction-diffusion equations and systems and integro-differential equations. We describe several counterintuitive results, where unfavorable factors for the persistence and spreading of a population tend to promote diversity in this population. In particular, we show that the Allee effect, the existence of a competitor species, as well as the presence of climate constraints are factors which can promote diversity during a colonization. We also show that long distance dispersal events lead to a higher diversity, whereas the existence of a nonreproductive juvenile stage does not affect the neutral diversity in a range-expanding population

Oscillatory periodic pattern dynamics in hyperbolic reaction-advection-diffusion models

In this work we consider a quite general class of two-species hyperbolic reaction-advection-diffusion system with the main aim of elucidating the role played by inertial effects in the dynamics of oscillatory periodic patterns. To this aim, first, we use linear stability analysis techniques to deduce the conditions under which wave (or oscillatory Turing) instability takes place. Then, we apply multiple-scale weakly nonlinear analysis to determine the equation which rules the spatiotemporal evolution of pattern amplitude close to criticality. This investigation leads to a cubic complex Ginzburg-Landau (CCGL) equation which, owing to the functional dependence of the coefficients here involved on the inertial times, reveals some intriguing consequences. To show in detail the richness of such a scenario, we present, as an illustrative example, the pattern dynamics occurring in the hyperbolic generalization of the extended Klausmeier model. This is a simple two-species model used to describe the migration of vegetation stripes along the hillslope of semiarid environments. By means of a thorough comparison between analytical predictions and numerical simulations, we show that inertia, apart from enlarging the region of the parameter plane where wave instability occurs, may also modulate the key features of the coherent structures, solution of the CCGL equation. In particular, it is proven that inertial effects play a role, not only during transient regime from the spatially-homogeneous steady state toward the patterned state, but also in altering the amplitude, the wavelength, the angular frequency, and even the stability of the phase-winding solutions

Partial Differential Equations in Ecology

Partial differential equations (PDEs) have been used in theoretical ecology research for more than eighty years. Nowadays, along with a variety of different mathematical techniques, they remain as an efficient, widely used modelling framework; as a matter of fact, the range of PDE applications has even become broader. This volume presents a collection of case studies where applications range from bacterial systems to population dynamics of human riots

Applications of stability theory to ecological problems

The goal of ecology is to investigate the interactions among organisms and their environment. However, ecological systems often exhibit complex dynamics. The application of mathematics to ecological problems has made tremendous progress over the years and many mathematical methods and tools have been developed for the exploration, whether analytical or numerical, of these dynamics. Mathematicians often study ecological systems by modelling them with partial differential equations (PDEs). Calculating the stability of solutions to these PDE systems is a classical question. This thesis first explores the concept of stability in the context of predator-prey invasions. Many ecological systems exhibit multi-year cycles. In such systems, invasions have a complicated spatiotemporal structure. In particular, it is common for unstable steady states to exist as long-term transients behind the invasion front, a phenomenon known as dynamical stabilisation. We combine absolute stability theory and computation to predict how the width of the stabilised region depends on parameter values. We develop our calculations in the context of a model for a cyclic predator-prey system, in which the invasion front and spatiotemporal oscillations of predators and prey are separated by a region in which the coexistence steady state is dynamically stabilised. Vegetation pattern formation in water-limited environments is another topic where stability theory plays a key role; indeed in mathematical models, these patterns are often the results of the dynamics that arise from perturbations to an unstable homogeneous steady state. Vegetation patterns are widespread in semi-deserts and aerial photographs of arid and semi-arid ecosystems have shown several kilometers square of these patterns. On hillsides in particular, vegetation is organised into banded spatial patterns. We first choose a domain in parameter space and calculate the boundary of the region in parameter space where pattern solutions exist. Finally we conclude with investigating how changes in the mean annual rainfall affect the properties of pattern solutions. Our work also highlights the importance of research on the calculation of the absolute spectrum for non-constant solutions

The stability of model ecosystems

Ecologists would like to understand how complexity persists in nature. In this thesis I have taken two fundamentally different routes to study ecosystem stability of model ecosystems: classical community ecology and classical population ecology. In community ecology models, we can study the mathematical mechanisms of stability in general, large model ecosystems. In population ecology models, fewer species are studied but greater detail of species interactions can be incorporated. Within these alternative contexts, this thesis contributes to two consuming issues concerning the stability of ecological systems: the ecosystem stability-complexity debate; and the causes of cyclic population dynamics. One of the major unresolved issues in community ecology is the relationship between ecosystem stability and complexity. In 1958 Charles Elton made the conjecture that the stability of an ecological system was coupled to its complexity and this could be a “wise principle of co-existence between man and nature” with which ecologists could argue the case for the conservation of nature for all species, including man. The earliest and simplest model systems were randomly constructed and exhibited a negative association between stability and complexity. This finding sparked the stability-complexity debate and initiated the search for organising principles that enhanced stability in real ecosystems. One of the universal laws of ecology is that ecosystems contain many rare and few common species. In this thesis, I present analytical arguments and numerical results to show that the stability of an ecosystem can increase with complexity when the abundance distribution is characterized by a skew towards many rare species. This work adds to the growing number of conditions under which the negative stability - complexity relationship can been inverted in theoretical studies. While there is growing evidence that the stability-complexity debate is progressing towards a resolution, community ecology has become increasingly subject to major criticism. A long-standing criticism is the reliance on local stability analysis. There is growing recognition that a global property called permanence is a more satisfactory definition of ecosystem stability because it tests only whether species can coexist. Here I identify and explain a positive correlation between the probability of local stability and permanence, which suggests local stability is a better measure of species coexistence than previously thought. While this offers some relief, remaining issues cause the stability-complexity debate to evade clear resolution and leave community ecology in a poor position to argue for the conservation of natural diversity for the benefit of all species. In classical population ecology, a major unresolved issue is the cause of non-equilibrium population dynamics. In this thesis, I use models to study the drivers of cyclic dynamics in Scottish populations of mountain hares (Lepus timidus), for the first time in this system. Field studies currently favour the hypothesis that parasitism by a nematode Trichostrongylus retortaeformis drives the hare cycles, and theory predicts that the interaction should induce cycling. Initially I used a simple, strategic host-parasite model parameterised using available empirical data to test the superficial concordance between theory and observation. I find that parasitism could not account for hare cycles. This verdict leaves three options: either the parameterisation was inadequate, there were missing important biological details or simply that parasites do not drive host cycles. Regarding the first option, reliable information for some hare-parasite model parameters was lacking. Using a rejection-sampling approach motivated by Bayesian methods, I identify the most likely parameter set to predict observed dynamics. The results imply that the current formulation of the hare-parasite model can only generate realistic dynamics when parasite effects are significantly larger than current empirical estimates, and I conclude it is likely that the model contains an inadequate level of detail. The simple strategic model was mathematically elegant and allowed mathematical concepts to be employed in analysis, but the model was biologically naïve. The second model is the antipode of the first, an individual based model (IBM) steeped in biological reality that can only be studied by simulation. Whilst most highly detailed tactical models are developed as a predictive tool, I instead structurally perturb the IBM to study the ecological processes that may drive population cycles in mountain hares. The model allows delayed responses to life history by linking maternal body size and parasite infection to the future survival and fecundity of offspring. By systematically removing model structure I show that these delayed life history effects are weakly destabilising and allow parameters to lie closer to empirical estimates to generate observed hare population cycles. In a third model I structurally modify the simple strategic host-parasite model to make it spatially explicit by including diffusion of mountain hares and corresponding advection of parasites (transportation with host). From initial simulations I show that the spatially extended host-parasite equations are able to generate periodic travelling waves (PTWs) of hare and parasite abundance. This is a newly documented behaviour in these widely used host-parasite equations. While PTWs are a new potential scenario under which cyclic hare dynamics could be explained, further mathematical development is required to determine whether adding space can generate realistic dynamics with parameters that lie closer to empirical estimates. In the general thesis discussion I deliberate on whether a hare-parasite model has been identified which can be considered the right balance between abstraction and relevant detail for this system

Rhythms and Evolution: Effects of Timing on Survival

The evolution of metabolism regulation is an intertwined process, where different strategies are constantly being developed towards a cognitive ability to perceive and respond to an environment. Organisms depend on an orchestration of a complex set of chemical reactions: maintaining homeostasis with a changing environment, while simultaneously sending material and energetic resources to where they are needed. The success of an organism requires efficient metabolic regulation, highlighting the connection between evolution, population dynamics and the underlying biochemistry. In this work, I represent organisms as coupled information-processing networks, that is, gene-regulatory networks receiving signals from the environment and acting on chemical reactions, eventually affecting material flows. I discuss the mechanisms through which metabolism control is improved during evolution and how the nonlinearities of competition influence this solution-searching process. The propagation of the populations through the resulting landscapes generally point to the role of the rhythm of cell division as an essential phenotypic feature driving evolution. Subsequently, as it naturally follows, different representations of organisms as oscillators are constructed to indicate more precisely how the interplay between competition, maturation timing and cell-division synchronisation affects the expected evolutionary outcomes, not always leading to the \"survival of the fastest\"

Spreading speeds and traveling waves in some population models.

Virtually every ecosystem has been invaded by exotic organisms with potentially drastic consequences for the native fauna or flora. Studying the forms and rates of invading species has been an important topic in spatial ecology. We investigate two two-species competition models with Allee effects in the forms of reaction-diffusion equations and integro-difference equations. We discuss the spatial transitions from a mono-culture equilibrium to a coexistence equilibrium or a different mono-culture equilibrium in these models. We provide formulas for the spreading speeds based on the linear determinacy and show the results on the existence of traveling waves. We also study a two-sex stage-structured model. We carry out initial analysis for the spreading speed and conduct numerical simulations on the traveling waves and spreading speeds in the two-sex model