Physiologically structured models - from versatile technique to ecological theory

A ubiquitous feature of natural communities is the variation in size that can be observed between organisms, a variation that to a substantial degree is intraspecific. Size variation within species by necessity implies that ecological interactions vary both in intensity and type over the life cycle of an individual. Physiologically structured population models (PSPMs) constitute a modelling approach especially designed to analyse these size-dependent interactions as they explicitly link individual level processes such as consumption and growth to population dynamics. We discuss two cases where PSPMs have been used to analyse the dynamics of size-structured populations. In the first case, a model of a size-structured consumer population feeding on a non-structured prey was successful in predicting both qualitative (mechanisms) and quantitative (individual growth, survival, cycle amplitude) aspects of the population dynamics of a planktivorous fish population. We conclude that single generation cycles as a result of intercohort competition is a general outcome of size-structured consumer–resource interactions. In the second case, involving both cohort competition and cannibalism, we show that PSPMs may predict double asymptotic growth trajectories with individuals ending up as giants. These growth trajectories, which have also been observed in field data, could not be predicted from individual level information, but are emergent properties of the population feedback on individual processes. In contrast to the size-structured consumer–resource model, the dynamics in this case cannot be reduced to simpler lumped stage-based models, but can only be analysed within the domain of PSPMs. Parameter values used in PSPMs adhere to the individual level and are derived independently from the system at focus, whereas model predictions involve both population level processes and individual level processes under conditions of population feedback. This leads to an increased ability to test model predictions but also to a larger set of variables that is predicted at both the individual and population level. The results turn out to be relatively robust to specific model assumptions and thus render a higher degree of generality than purely individual-based models. At the same time, PSPMs offer a much higher degree of realism, precision and testing ability than lumped stage-based or non-structured models. The results of our analyses so far suggest that also in more complex species configurations only a limited set of mechanisms determines the dynamics of PSPMs. We therefore conclude that there is a high potential for developing an individual-based, size-dependent community theory using PSPMs.