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We consider a hierarchically structured population in which the amount of resources an individual has access to is affected by individuals that are larger, and that the intake of resources by an individual only affects directly the growth rate of the individual. We formulate a deterministic model, which takes the form of a delay equation for the population birth rate. We also formulate an individual based stochastic model, and study the relationship between the two models. In particular the stationary birth rate of the deterministic model is compared to that of the quasi-stationary birth rate of the stochastic model. Since the quasi-stationary birth rate cannot be obtained explicitly, we derive a formula to approximate it. We show that the stationary birth rate of the deterministic model can be obtained as the large population limit of the quasi-stationary birth rate of the stochastic model. This relation suggests that the deterministic model is a good approximation of the stochastic model when the number of individuals is sufficiently large., Comment: 3 figures, to appear in Journal of Applied Mathematics and Computing

In this paper we investigate a structured population model with distributed delay. Our model incorporates two different types of nonlinearities. Specifically we assume that individual growth and mortality are affected by scramble competition, while fertility is affected by contest competition. In particular, we assume that there is a hierarchical structure in the population, which affects mating success. The dynamical behavior of the model is analysed via linearisation by means of semigroup and spectral methods. In particular, we introduce a reproduction function and use it to derive linear stability criteria for our model. Further we present numerical simulations to underpin the stability results we obtained., Comment: 4 figures

We consider a population organised hierarchically with respect to size in such a way that the growth rate of each individual depends only on the presence of larger individuals. As a concrete example one might think of a forest, in which the incidence of light on a tree (and hence how fast it grows) is affected by shading of taller trees. The model is formulated as a delay equation, more specifically a scalar renewal equation, for the population birth rate. After discussing the well-posedness of the model, we analyse how many stationary birth rates the equation can have in terms of the functional parameters of the model. In particular we show that, under reasonable and rather general assumptions, only one stationary birth rate can exist besides the trivial one (associated to the state in which there are no individuals and the population birth rate is zero). We give conditions for this non-trivial stationary birth rate to exist and we analyse its stability using the principle of linearised stability for delay equations. Finally we relate the results to an alternative formulation of the model taking the form of a quasilinear partial differential equation for the population size-density.

To describe the dynamics of a size-structured population and its unstructured resource, we formulate bookkeeping equations in two different ways. The first, called the PDE formulation, is rather standard. It employs a first order partial differential equation, with a non-local boundary condition, for the size-density of the consumer, coupled to an ordinary differential equation for the resource concentration. The second is called the DELAY formulation and employs a renewal equation for the population level birth rate of the consumer, coupled to a delay differential equation for the (history of the) resource concentration. With each of the two formulations we associate a constructively defined semigroup of nonlinear solution operators. The two semigroups are intertwined by a non-invertible operator. In this paper we delineate in what sense the two semigroups are equivalent. In particular, we (i) identify conditions on both the model ingredients and the choice of state space that guarantee that the intertwining operator is surjective, (ii) focus on large time behaviour and (iii) consider full orbits, i.e., orbits defined for time running from $-\infty $ to $+\infty $. Conceptually, the PDE formulation is by far the most natural one. It has, however, the technical drawback that the solution operators are not differentiable, precluding rigorous linearisation. For the delay formulation, one can (under certain conditions concerning the model ingredients) prove the differentiability of the solution operators and establish the Principle of Linearised Stability. Next the equivalence of the two formulations yields a rather indirect proof of this principle for the PDE formulation., Comment: To appear in Mathematical Models and Methods in the Applied Sciences

According to the World Health Organization the global incidence rate of dengue infections have risen drastically in recent years. It is estimated that globally the number of new infections is in the range of $100$ to $400$ million per annum. At the same time a number of recent studies reported the existence of Wolbachia strains, which inhibit dengue virus replication in mosquito species that are primary vectors for dengue transmission. In this study we focus on the impact of Wolbachia on dengue transmission dynamics in an age-structured human population. We introduce a mathematical model, which takes into account age-related effects, such as age-dependent human recovery and mortality rates, as well as age-dependent vector to human dengue transmission efficacy. We deduce an explicit formula for the basic reproduction number $\mathcal{R}_0$, which allows us to assess the impact of the above mentioned age-related effects on the local asymptotic stability of the dengue free equilibrium. The formula we deduce for $\mathcal{R}_0$ also shows the intricate relationship between human demography and the presence of a dengue inhibiting Wolbachia strain.

We introduce a system of differential equations to assess the impact of (self-)quarantine of symptomatic infectious individuals on disease dynamics. To this end we depart from using the classic bilinear infection process, but remain still within the framework of the mass-action assumption. From the mathematical point of view our model is interesting due to the lack of continuous differentiability at disease free steady states, which implies also that the basic reproductive number cannot be computed following established approaches for certain parameter values. However, we parametrise our mathematical model using published values from the COVID-19 literature, and analyse the model simulations. We also contrast model simulations against publicly available COVID-19 test data focusing on the first wave of the pandemic during March - July 2020 in the UK. Our simulations indicate that actual peak case numbers might have been as much as 200 times higher than the reported positive test cases during the first wave in the UK. We find that very strong adherence to self-quarantine rules yields (only) a reduction of 22$\%$ of peak numbers and delays the onset of the peak by approximately 30-35 days. However, during the early phase of the outbreak the impact of (self)-quarantine is much more significant. We also take into account the effect of a national lockdown in a simplistic way by reducing the effective susceptible population size. We find that in case of a 90$\%$ reduction of the effective susceptible population size, strong adherence to self-quarantine still only yields a 25$\%$ reduction of peak infectious numbers when compared to low adherence. This is due to the significant number of asymptomatic infectious individuals in the population.

In this work we establish conditions which guarantee the existence of (strictly) positive steady states of a nonlinear structured population model. In our framework the steady state formulation amounts to recasting the nonlinear problem as a family of eigenvalue problems combined with a fixed point problem. Amongst other things our formulation requires us to control the growth behaviour of the spectral bound of a family of linear operators along positive rays. For the specific class of model we consider here, this presents a considerable challenge. We are going to show that the spectral bound of the family of operators, arising from the steady state formulation, can be controlled by perturbations in the domain of the generators (only). These new boundary perturbation results are particularly important for models exhibiting fertility controlled dynamics. As an important by-product of the application of the boundary perturbation results we employ here we recover (using a recent theorem by H.R. Thieme) the familiar net reproduction number (or function) for models with single state at birth, which include for example the classic McKendrick (linear) and Gurtin-McCamy (non-linear) age-structured models.

In this paper we consider a physiologically structured population model with distributed states at birth, formulated on the space of non-negative Radon measures. Using a characterisation of the pre-dual space of bounded Lipschitz functions, we show how to apply the theory of strongly continuous positive semigroups to such a model. In particular, we establish the exponential convergence of solutions to a one-dimensional global attractor.

The goal of this note is to present a general approach to define the net reproduction function for a large class of nonlinear physiologically structured population models. In particular, we are going to show that this can be achieved in a natural way by reformulating a nonlinear problem as a family of linear ones; each of the linear problems describing the evolution of the population in a different, but constant environment. The reformulation of a nonlinear population model as a family of linear ones is a new approach, and provides an elegant way to study qualitative questions, for example the existence of positive steady states. To define the net reproduction number for any fixed (constant) environment, i.e. for the linear models, we use a fairly recent spectral theoretic result, which characterizes the connection between the spectral bound of an unbounded operator and the spectral radius of a corresponding bounded operator. For nonlinear models, varying the environment naturally leads to a net reproduction function., Comment: To appear in Mathematical Modelling of Natural Phenomena

We investigate a partial differential equation model of a cancer cell population, which is structured with respect to age and telomere length of cells. We assume a continuous telomere length structure, which is applicable to the clonal evolution model of cancer cell growth. This model has a non-standard non-local boundary condition. We establish global existence of solutions and study their qualitative behaviour. We study the effect of telomere restoration on cancer cell dynamics. Our results indicate that without telomere restoration, the cell population extinguishes. With telomere restoration, exponential growth occurs in the linear model. We further characterise the specific growth behaviour of the cell population for special cases. We also study the effects of crowding induced mortality on the qualitative behaviour, and the existence and stability of steady states of a nonlinear model incorporating crowding effect. We present examples and extensive numerical simulations, which illustrate the rich dynamic behaviour of the linear and nonlinear models., Comment: 36 pages, 12 figures

In this work first we consider a physiologically structured population model with a distributed recruitment process. That is, our model allows newly recruited individuals to enter the population at all possible individual states, in principle. The model can be naturally formulated as a first order partial integro-differential equation, and it has been studied extensively. In particular, it is well-posed on the biologically relevant state space of Lebesgue integrable functions. We also formulate a delayed integral equation (renewal equation) for the distributed birth rate of the population. We aim to illustrate the connection between the partial integro-differential and the delayed integral equation formulation of the model utilising a recent spectral theoretic result. In particular, we consider the equivalence of the steady state problems in the two different formulations, which then leads us to characterise irreducibility of the semigroup governing the linear partial integro-differential equation. Furthermore, using the method of characteristics, we investigate the connection between the time dependent problems. In particular, we prove that any (non-negative) solution of the delayed integral equation determines a (non-negative) solution of the partial differential equation and vice versa. The results obtained for the particular distributed states at birth model then lead us to present some very general results, which establish the equivalence between a general class of partial differential and delay equation, modelling physiologically structured populations., Comment: 28 pages, to appear in Mathematical Methods in the Applied Sciences

We introduce and investigate an SIS-type model for the spread of an infectious disease, where the infected population is structured with respect to the different strain of the virus/bacteria they are carrying. Our aim is to capture the interesting scenario when individuals infected with different strains cause secondary (new) infections at different rates. Therefore, we consider a nonlinear infection process, which generalises the bilinear process arising from the classic mass-action assumption. Our main motivation is to study competition between different strains of a virus/bacteria. From the mathematical point of view, we are interested whether the nonlinear infection process leads to a well-posed model. We use a semilinear formulation to show global existence and positivity of solutions up to a critical value of the exponent in the nonlinearity. Furthermore, we establish the existence of the endemic steady state for particular classes of nonlinearities., Comment: 21 pages, to appear in Nonlinear Analysis: Real World Applications

Wolbachia is possibly the most studied reproductive parasite of arthropod species. It appears to be a promising candidate for biocontrol of some mosquito borne diseases. We begin by developing a sex-structured model for a Wolbachia infected mosquito population. Our model incorporates the key effects of Wolbachia infection including cytoplasmic incompatibility and male killing. We also allow the possibility of reduced reproductive output, incomplete maternal transmission, and different mortality rates for uninfected/infected male/female individuals. We study the existence and local stability of equilibria, including the biologically relevant and interesting boundary equilibria. For some biologically relevant parameter regimes there may be multiple coexistence steady states including, very importantly, a coexistence steady state in which Wolbachia infected individuals dominate. We also extend the model to incorporate West Nile virus (WNv) dynamics, using an SEI modelling approach. Recent evidence suggests that a particular strain of Wolbachia infection significantly reduces WNv replication in Aedes aegypti. We model this via increased time spent in the WNv-exposed compartment for Wolbachia infected female mosquitoes. A basic reproduction number $R_0$ is computed for the WNv infection. Our results suggest that, if the mosquito population consists mainly of Wolbachia infected individuals, WNv eradication is likely if WNv replication in Wolbachia infected individuals is sufficiently reduced., Comment: to appear in Journal of Mathematical Biology

We employ partial integro-differential equations to model trophic interaction in a spatially extended heterogeneous environment. Compared to classical reaction-diffusion models, this framework allows us to more realistically describe the situation where movement of individuals occurs on a faster time scale than the demographic (population) time scale, and we cannot determine population growth based on local density. However, most of the results reported so far for such systems have only been verified numerically and for a particular choice of model functions, which obviously casts doubts about these findings. In this paper, we analyse a class of integro-differential predator-prey models with a highly mobile predator in a heterogeneous environment, and we reveal the main factors stabilizing such systems. In particular, we explore an ecologically relevant case of interactions in a highly eutrophic environment, where the prey carrying capacity can be formally set to 'infinity'. We investigate two main scenarios: (i) the spatial gradient of the growth rate is due to abiotic factors only, and (ii) the local growth rate depends on the global density distribution across the environment (e.g. due to non-local self-shading). For an arbitrary spatial gradient of the prey growth rate, we analytically investigate the possibility of the predator-prey equilibrium in such systems and we explore the conditions of stability of this equilibrium. In particular, we demonstrate that for a Holling type I (linear) functional response, the predator can stabilize the system at low prey density even for an 'unlimited' carrying capacity. We conclude that the interplay between spatial heterogeneity in the prey growth and fast displacement of the predator across the habitat works as an efficient stabilizing mechanism., Comment: 2 figures; appendices available on request. To appear in the Bulletin of Mathematical Biology

In this paper we explore the eco-evolutionary dynamics of a predator-prey model, where the prey population is structured according to a certain life history trait. The trait distribution within the prey population is the result of interplay between genetic inheritance and mutation, as well as selectivity in the consumption of prey by the predator. The evolutionary processes are considered to take place on the same time scale as ecological dynamics, i.e. we consider the evolution to be rapid. Previously published results show that population structuring and rapid evolution in such predator-prey system can stabilise an otherwise globally unstable dynamics even with an unlimited carrying capacity of prey. However, those findings were only based on direct numerical simulation of equations and obtained for particular parametrisations of model functions, which obviously calls into question the correctness and generality of the previous results. The main objective of the current study is to treat the model analytically and consider various parametrisations of predator selectivity and inheritance kernel. We investigate the existence of a coexistence stationary state in the model and carry out stability analysis of this state. We derive expressions for the Hopf bifurcation curve which can be used for constructing bifurcation diagrams in the parameter space without the need for a direct numerical simulation of the underlying integro-differential equations. We analytically show the possibility of stabilisation of a globally unstable predator-prey system with prey structuring. We prove that the coexistence stationary state is stable when the saturation in the predation term is low. Finally, for a class of kernels describing genetic inheritance and mutation we show that stability of the predator-prey interaction will require a selectivity of predation according to the life trait., Comment: 28 pages, 1 figure, to appear in MMNP

We study the question of existence of positive steady states of nonlinear evolution equations. We recast the steady state equation in the form of eigenvalue problems for a parametrised family of unbounded linear operators, which are generators of strongly continuous semigroups; and a fixed point problem. In case of irreducible governing semigroups we consider evolution equations with non-monotone nonlinearities of dimension two, and we establish a new fixed point theorem for set-valued maps. In case of reducible governing semigroups we establish results for monotone nonlinearities of any finite dimension $n$. In addition, we establish a non-quasinilpotency result for a class of strictly positive operators, which are neither irreducible nor compact, in general. We illustrate our theoretical results with examples of partial differential equations arising in structured population dynamics. In particular, we establish existence of positive steady states of a size-structured juvenile-adult and a structured consumer-resource population model, as well as for a selection-mutation model with distributed recruitment process., Comment: to appear in SIAM Journal on Mathematical Analysis

We consider a nonlinear structured population model with a distributed recruitment term. The question of the existence of non-trivial steady states can be treated (at least!) in three different ways. One approach is to study spectral properties of a parametrized family of unbounded operators. The alternative approach, on which we focus here, is based on the reformulation of the problem as an integral equation. In this context we introduce a density dependent net reproduction rate and discuss its relationship to a biologically meaningful quantity. Finally, we briefly discuss a third approach, which is based on the finite rank approximation of the recruitment operator., Comment: To appear in Computers and Mathematics with Applications

We introduce a nonlinear structured population model with diffusion in the state space. Individuals are structured with respect to a continuous variable which represents a pathogen load. The class of uninfected individuals constitutes a special compartment that carries mass, hence the model is equipped with generalized Wentzell (or dynamic) boundary conditions. Our model is intended to describe the spread of infection of a vertically transmitted disease, for example Wolbachia in a mosquito population. Therefore the (infinite dimensional) nonlinearity arises in the recruitment term. First we establish global existence of solutions and the Principle of Linearised Stability for our model. Then, in our main result, we formulate simple conditions, which guarantee the existence of non-trivial steady states of the model. Our method utilizes an operator theoretic framework combined with a fixed point approach. Finally, in the last section we establish a sufficient condition for the local asymptotic stability of the positive steady state.

We propose a hybrid dynamical system approach to model the evolution of a pathogen that experiences different selective pressures according to a stochastic process. In every environment, the evolution of the pathogen is described by a version of the Fisher-Haldane-Wright equation while the switching between environments follows a Markov jump process. We investigate how the qualitative behavior of a simple single-host deterministic system changes when the stochastic switching process is added. In particular, we study the stability in probability of monomorphic equilibria. We prove that in a "constantly" fluctuating environment, the genotype with the highest mean fitness is asymptotically stable in probability. However, if the probability of host switching depends on the genotype composition of the population, polymorphism can be stably maintained. This is a corrected version of the paper that appeared in Mathematical Biosciences 240 (2012), p. 70-75. A corrigendum has appeared in the same journal., Comment: 15 pages, 4 figures

We consider a class of physiologically structured population models, a first order nonlinear partial differential equation equipped with a nonlocal boundary condition, with a constant external inflow of individuals. We prove that the linearised system is governed by a quasicontraction semigroup. We also establish that linear stability of equilibrium solutions is governed by a generalized net reproduction function. In a special case of the model ingredients we discuss the nonlinear dynamics of the system when the spectral bound of the linearised operator equals zero, i.e. when linearisation does not decide stability. This allows us to demonstrate, through a concrete example, how immigration might be beneficial to the population. In particular, we show that from a nonlinearly unstable positive equilibrium a linearly stable and unstable pair of equilibria bifurcates. In fact, the linearised system exhibits bistability, for a certain range of values of the external inflow, induced potentially by All\'{e}e-effect., Comment: to appear in Journal of Applied Mathematics and Computing

We introduce and investigate a series of models for an infection of a diplodiploid host species by the bacterial endosymbiont \textit{Wolbachia}. The continuous models are characterized by partial vertical transmission, cytoplasmic incompatibility and fitness costs associated with the infection. A particular aspect of interest is competitions between mutually incompatible strains. We further introduce an age-structured model that takes into account different fertility and mortality rates at different stages of the life cycle of the individuals. With only a few parameters, the ordinary differential equation models exhibit already interesting dynamics and can be used to predict criteria under which a strain of bacteria is able to invade a population. Interestingly, but not surprisingly, the age-structured model shows significant differences concerning the existence and stability of equilibrium solutions compared to the unstructured model., Comment: 24 pages, 6 figures, revised and expanded

In this work we introduce and analyze a linear size-structured population model with infinite states-at-birth. We model the dynamics of a population in which individuals have two distinct life-stages: an "active" phase when individuals grow, reproduce and die and a second "resting" phase when individuals only grow. Transition between these two phases depends on individuals' size. First we show that the problem is governed by a positive quasicontractive semigroup on the biologically relevant state space. Then we investigate, in the framework of the spectral theory of linear operators, the asymptotic behavior of solutions of the model. We prove that the associated semigroup has, under biologically plausible assumptions, the property of asynchronous exponential growth., Comment: 1 figure, to appear in Positivity

Motivated by structured parasite populations in aquaculture we consider a class of size-structured population models, where individuals may be recruited into the population with distributed states at birth. The mathematical model which describes the evolution of such a population is a first-order nonlinear partial integro-differential equation of hyperbolic type. First, we use positive perturbation arguments and utilise results from the spectral theory of semigroups to establish conditions for the existence of a positive equilibrium solution of our model. Then, we formulate conditions that guarantee that the linearised system is governed by a positive quasicontraction semigroup on the biologically relevant state space. We also show that the governing linear semigroup is eventually compact, hence growth properties of the semigroup are determined by the spectrum of its generator. In the case of a separable fertility function, we deduce a characteristic equation, and investigate the stability of equilibrium solutions in the general case using positive perturbation arguments., Comment: to appear in Mathematical Modelling of Natural Phenomena

In this work we consider a size-structured cannibalism model with the model ingredients (fertility, growth, and mortality rate) depending on size (ranging over an infinite domain) and on a general function of the standing population (environmental feedback). Our focus is on the asymptotic behavior of the system, in particular on the effect of cannibalism on the long-term dynamics. To this end, we formally linearize the system about steady state and establish conditions in terms of the model ingredients which yield uniform exponential stability of the governing linear semigroup. We also show how the point spectrum of the linearized semigroup generator can be characterized in the special case of a separable attack rate and establish a general instability result. Further spectral analysis allows us to give conditions for asynchronous exponential growth of the linear semigroup.

In the present paper we analyze the linear stability of a hierarchical size-structured population model where the vital rates (mortality, fertility and growth rate) depend both on size and a general functional of the population density ("environment"). We derive regularity properties of the governing linear semigroup, implying that linear stability is governed by a dominant real eigenvalue of the semigroup generator, which arises as a zero of an associated characteristic function. In the special case where neither the growth rate nor the mortality depend on the environment, we explicitly calculate the characteristic function and use it to formulate simple conditions for the linear stability of population equilibria. In the general case we derive a dissipativity condition for the linear semigroup, thereby characterizing exponential stability of the steady state.