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We design a linear chain trick algorithm for dynamical systems for which we have oscillatory time histories in the distributed time delay. We make use of this algorithmic framework to analyse memory effects in disease evolution in a population. The modelling is based on a susceptible-infected-recovered SIR-model and on a susceptible-exposed-infected-recovered SEIR-model through a kernel that dampens the activity based on the recent history of infectious individuals. This corresponds to adaptive behavior in the population or through governmental non-pharmaceutical interventions. We use the linear chain trick to show that such a model may be written in a Markovian way, and we analyze the stability of the system. We find that the adaptive behavior gives rise to either a stable equilibrium point or a stable limit cycle for a close to constant number of susceptibles, i.e. locally in time. We also show that the attack rate for this model is lower than it would be without the dampening, although the adaptive behavior disappears as time goes to infinity and the number of infected goes to zero.

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The severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) has been spreading worldwide for over two years, with millions of reported cases and deaths. The deployment of mathematical modeling in the fight against COVID-19 has recorded tremendous success. However, most of these models target the epidemic phase of the disease. The development of safe and effective vaccines against SARS-CoV-2 brought hope of safe reopening of schools and businesses and return to pre-COVID normalcy, until mutant strains like the Delta and Omicron variants, which are more infectious, emerged. A few months into the pandemic, reports of the possibility of both vaccine- and infection-induced immunity waning emerged, thereby indicating that COVID-19 may be with us for longer than earlier thought. As a result, to better understand the dynamics of COVID-19, it is essential to study the disease with an endemic model. In this regard, we developed and analyzed an endemic model of COVID-19 that incorporates the waning of both vaccine- and infection-induced immunities using distributed delay equations. Our modeling framework assumes that the waning of both immunities occurs gradually over time at the population level. We derived a nonlinear ODE system from the distributed delay model and showed that the model could exhibit either a forward or backward bifurcation depending on the immunity waning rates. Having a backward bifurcation implies that $ R_c < 1 $ is not sufficient to guarantee disease eradication, and that the immunity waning rates are critical factors in eradicating COVID-19. Our numerical simulations show that vaccinating a high percentage of the population with a safe and moderately effective vaccine could help in eradicating COVID-19.

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We introduce a distributed-delay differential equation disease spread model for COVID-19 spread. The model explicitly incorporates the population's time-dependent vaccine uptake and incorporates a gamma-distributed temporary immunity period for both vaccination and previous infection. We validate the model on COVID-19 cases and deaths data from the state of Michigan and use the calibrated model to forecast the spread and impact of the disease under a variety of realistic booster vaccine strategies. The model suggests that the mean immunity duration for individuals after vaccination is 350 days and after a prior infection is 242 days. Simulations suggest that both high population-wide adherence to vaccination mandates and a more-than-annually frequency of booster doses will be required to contain outbreaks in the future.

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Gamma distributed delay differential equations (DDEs) arise naturally in many modelling applications. However, appropriate numerical methods for generic gamma distributed DDEs have not previously been implemented. Modellers have therefore resorted to approximating the gamma distribution with an Erlang distribution and using the linear chain technique to derive an equivalent system of ordinary differential equations (ODEs). In this work, we address the lack of appropriate numerical tools for gamma distributed DDEs in two ways. First, we develop a functional continuous Runge-Kutta (FCRK) method to numerically integrate the gamma distributed DDE without resorting to Erlang approximation. We prove the fourth-order convergence of the FCRK method and perform numerical tests to demonstrate the accuracy of the new numerical method. Nevertheless, FCRK methods for infinite delay DDEs are not widely available in existing scientific software packages. As an alternative approach to solving gamma distributed DDEs, we also derive a hypoexponential approximation of the gamma distributed DDE. This hypoexponential approach is a more accurate approximation of the true gamma distributed DDE than the common Erlang approximation but, like the Erlang approximation, can be formulated as a system of ODEs and solved numerically using standard ODE software. Using our FCRK method to provide reference solutions, we show that the common Erlang approximation may produce solutions that are qualitatively different from the underlying gamma distributed DDE. However, the proposed hypoexponential approximations do not have this limitation. Finally, we apply our hypoexponential approximations to perform statistical inference on synthetic epidemiological data to illustrate the utility of the hypoexponential approximation.

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To mitigate the harmful effects of the COVID-19 pandemic, world countries have resorted - though with different timing and intensities - to a range of interventions. These interventions and their relaxation have shaped the epidemic into a multi-phase form, namely an early invasion phase often followed by a lockdown phase, whose unlocking triggered a second epidemic wave, and so on. In this article, we provide a kinematic description of an epidemic whose time course is subdivided by mitigation interventions into a sequence of phases, on the assumption that interventions are effective enough to prevent the susceptible proportion to largely depart from 100% (or from any other relevant level). By applying this hypothesis to a general SIR epidemic model with age-since-infection and piece-wise constant contact and recovery rates, we supply a unified treatment of this multi-phase epidemic showing how the different phases unfold over time. Subsequently, by exploiting a wide class of infectiousness and recovery kernels allowing reducibility (either to ordinary or delayed differential equations), we investigate in depth a low-dimensional case allowing a non-trivial full analytical treatment also of the transient dynamics connecting the different phases of the epidemic. Finally, we illustrate our theoretical results by a fit to the overall Italian COVID-19 epidemic since March 2020 till February 2021 i.e., before the mass vaccination campaign. This show the abilities of the proposed model in effectively describing the entire course of an observed multi-phasic epidemic with a minimal set of data and parameters, and in providing useful insight on a number of aspects including e.g., the inertial phenomena surrounding the switch between different phases.

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The well known linear chain trick (LCT) allows modellers to derive mean field ODEs that assume gamma (Erlang) distributed passage times, by transitioning individuals sequentially through a chain of sub-states. The time spent in these sub-states is the sum of k exponentially distributed random variables, and is thus gamma distributed. The generalized linear chain trick (GLCT) extends this technique to the broader phase-type family of distributions, which includes exponential, Erlang, hypoexponential, and Coxian distributions. Phase-type distributions are the family of matrix exponential distributions on [0,∞) that represent the absorption time distributions for finite-state, continuous time Markov chains (CTMCs). Here we review CTMCs and phase-type distributions, then illustrate how to use the GLCT to efficiently build ODE models from underlying stochastic model assumptions. We introduce two novel model families by using the GLCT to generalize the Rosenzweig-MacArthur predator-prey model, and the SEIR model. We illustrate the kinds of complexity that can be captured by such models through multiple examples. We also show the benefits of using a GLCT-based model formulation to speed up the computation of numerical solutions to such models. These results highlight the intuitive nature, and utility, of using the GLCT to derive ODE models from first principles.

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A prototype SIR model with vaccination at birth is analyzed in terms of the stability of its endemic equilibrium. The information available on the disease influences the parents' decision on whether vaccinate or not. This information is modeled with a delay according to the Erlang distribution. The latter includes the degenerate case of fading memory as well as the limiting case of concentrated memory. The linear chain trick is the essential tool used to investigate the general case. Besides its novel analysis and that of the concentrated case, it is showed that through the linear chain trick a distributed delay approaches a discrete delay at a linear rate. A rigorous proof is given in terms of the eigenvalues of the associated linearized problems and extension to general models is also provided. The work is completed with several computations and relevant experimental results.

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In a physiologically structured population model (PSPM) individuals are characterised by continuous variables, like age and size, collectively called their i-state. The world in which these individuals live is characterised by another set of variables, collectively called the environmental condition. The model consists of submodels for (i) the dynamics of the i-state, e.g. growth and maturation, (ii) survival, (iii) reproduction, with the relevant rates described as a function of (i-state, environmental condition), (iv) functions of (i-state, environmental condition), like biomass or feeding rate, that integrated over the i-state distribution together produce the output of the population model. When the environmental condition is treated as a given function of time (input), the population model becomes linear in the state. Density dependence and interaction with other populations is captured by feedback via a shared environment, i.e., by letting the environmental condition be influenced by the populations' outputs. This yields a systematic methodology for formulating community models by coupling nonlinear input-output relations defined by state-linear population models. For some combinations of submodels an (infinite dimensional) PSPM can without loss of relevant information be replaced by a finite dimensional ODE. We then call the model ODE-reducible. The present paper provides (a) a test for checking whether a PSPM is ODE reducible, and (b) a catalogue of all possible ODE-reducible models given certain restrictions, to wit: (i) the i-state dynamics is deterministic, (ii) the i-state space is one-dimensional, (iii) the birth rate can be written as a finite sum of environment-dependent distributions over the birth states weighted by environment independent 'population outputs'. So under these restrictions our conditions for ODE-reducibility are not only sufficient but in fact necessary. Restriction (iii) has the desirable effect that it guarantees that the population trajectories are after a while fully determined by the solution of the ODE so that the latter gives a complete picture of the dynamics of the population and not just of its outputs.

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In this paper we generalize the Linear Chain Trick (LCT; aka the Gamma Chain Trick) to help provide modelers more flexibility to incorporate appropriate dwell time assumptions into mean field ODEs, and help clarify connections between individual-level stochastic model assumptions and the structure of corresponding mean field ODEs. The LCT is a technique used to construct mean field ODE models from continuous-time stochastic state transition models where the time an individual spends in a given state (i.e., the dwell time) is Erlang distributed (i.e., gamma distributed with integer shape parameter). Despite the LCT's widespread use, we lack general theory to facilitate the easy application of this technique, especially for complex models. Modelers must therefore choose between constructing ODE models using heuristics with oversimplified dwell time assumptions, using time consuming derivations from first principles, or to instead use non-ODE models (like integro-differential or delay differential equations) which can be cumbersome to derive and analyze. Here, we provide analytical results that enable modelers to more efficiently construct ODE models using the LCT or related extensions. Specifically, we provide (1) novel LCT extensions for various scenarios found in applications, including conditional dwell time distributions; (2) formulations of these LCT extensions that bypass the need to derive ODEs from integral equations; and (3) a novel Generalized Linear Chain Trick (GLCT) framework that extends the LCT to a much broader set of possible dwell time distribution assumptions, including the flexible phase-type distributions which can approximate distributions on [Formula: see text] and can be fit to data.

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