Fixation and extinction in time-fluctuating spatially structured metapopulations

Abstract

Bacteria evolve in volatile environments and complex spatial structures. Migration, fluctuations and environmental variability therefore have a significant impact on the evolution of microbial populations. Here, we consider a class of spatially explicit metapopulation models arranged as regular (circulation) graphs where wild-type and mutant cells compete in a time-fluctuating environment in which demes (subpopulations) are connected by slow cell migration. The carrying capacity is the same at each deme and endlessly switches between two values associated to harsh and mild environmental conditions. It is known that environmental variability can lead to population bottle-necks, following which the population is prone to fluctuation-induced extinction. Here, we analyse how slow migration, spatial structure, and fluctuations affect the phenomena of fixation and extinction on clique, cycle, and square lattice metapopulations. When the carrying capacity remains large, bottlenecks are weak and deme extinction can be ignored. The dynamics is thus captured by a coarse-grained description within which the probability and mean time of fixation are obtained analytically. This allows us to show that, in contrast to what happens in static environments, the mutant fixation probability depends on the rate of migration. We also show that the fixation probability and mean fixation time can exhibit a non-monotonic dependence on the switching rate. When the carrying capacity is small under harsh conditions, bottlenecks are strong, and the metapopulation evolution is shaped by the coupling of deme extinction and strain competition. This yields rich dynamical scenarios, among which we identify the best conditions to eradicate mutants without dooming the metapopulation to extinction. We offer an interpretation of these findings in the context of an idealised treatment strategy and discuss possible generalisations of our models.