Abstract

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.

Highlights

  • From the very beginning of community modelling, ordinary differential equations (ODEs) have been its main tool

  • The key question addressed in this paper is : under what conditions on the model ingredients is it possible to obtain the same input–output relation when the physiologically structured population model (PSPM) is replaced by a finite dimensional ODE? (This representation may have an interpretation in its own right, but this is not required.) If such a representation is possible, we say that the population model is reduced to the ODE or that the input–output relation is realised by it

  • Our focus should be on the biological problem: when is a physiologically structured model ODE representable? That is the question that matters, both practically and foundationally, for community modelling

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Summary

Introduction

From the very beginning of community modelling, ordinary differential equations (ODEs) have been its main tool This notwithstanding the fact that much earlier Euler (1760) and other mathematicians working on population dynamics had already considered age structured models, see (Bacaër 2008, 2011; Gyllenberg 2007) for more information on the history of population dynamics. For the practically important subset of cases where the population birth rate figures on the list of population outputs and with a single state variable on the level of the individuals, the last author solved this problem on a heuristic level already in 1989 during a holiday week in summer spent at the Department of Applied Physics of the University of Strathclyde An allusion to this was given in a “note added in print” to the paper (Metz and Diekmann 1991). The much shorter paper (Diekmann et al 2019) provides additional examples and may serve as a more friendly user guide to ODE-reducibility of structured population models

Biological context
The mathematical question
A tool
Testing combinations of model ingredients for ODE reducibility
A catalogue of ODE reducible models
For mathematicians: the context of justification
Physiologically structured population models
General considerations
A catalogue of models that admit a finite dimensional state representation
Some spadework
The model family F3
Applications to age- and stage-structured population models
Discussion
Full Text
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