Cooperation in Microbial Populations: Theory and Experimental Model Systems

Cooperative behavior, the costly provision of benefits to others, is common across all domains of life. This review article discusses cooperative behavior in the microbial world, mediated by the exchange of extracellular products called public goods. We focus on model species for which the production of a public good and the related growth disadvantage for the producing cells are well described. To unveil the biological and ecological factors promoting the emergence and stability of cooperative traits we take an interdisciplinary perspective and review insights gained from both mathematical models and well-controlled experimental model systems. Ecologically, we include crucial aspects of the microbial life cycle into our analysis and particularly consider population structures where an ensemble of local communities (sub populations) continuously emerge, grow, and disappear again. Biologically, we explicitly consider the synthesis and regulation of public good production. The discussion of the theoretical approaches includes general evolutionary concepts, population dynamics, and evolutionary game theory. As a specific but generic biological example we consider populations of Pseudomonas putida and its regulation and utilization of pyoverdines, iron scavenging molecules. The review closes with an overview on cooperation in spatially extended systems and also provides a critical assessment of the insights gained from the experimental and theoretical studies discussed. Current challenges and important new research opportunities are discussed, including the biochemical regulation of public goods, more realistic ecological scenarios resembling native environments, cell to cell signalling, and multi-species communities.Comment: Review article, 88 pages, 14 figure

Evolutionary game theory in growing populations

Existing theoretical models of evolution focus on the relative fitness advantages of different mutants in a population while the dynamic behavior of the population size is mostly left unconsidered. We here present a generic stochastic model which combines the growth dynamics of the population and its internal evolution. Our model thereby accounts for the fact that both evolutionary and growth dynamics are based on individual reproduction events and hence are highly coupled and stochastic in nature. We exemplify our approach by studying the dilemma of cooperation in growing populations and show that genuinely stochastic events can ease the dilemma by leading to a transient but robust increase in cooperationComment: 4 pages, 2 figures and 2 pages supplementary informatio

Range expansion with mutation and selection: dynamical phase transition in a two-species Eden model

The colonization of unoccupied territory by invading species, known as range expansion, is a spatially heterogeneous non-equilibrium growth process. We introduce a two-species Eden growth model to analyze the interplay between uni-directional (irreversible) mutations and selection at the expanding front. While the evolutionary dynamics leads to coalescence of both wild-type and mutant clusters, the non-homogeneous advance of the colony results in a rough front. We show that roughening and domain dynamics are strongly coupled, resulting in qualitatively altered bulk and front properties. For beneficial mutations the front is quickly taken over by mutants and growth proceeds Eden-like. In contrast, if mutants grow slower than wild-types, there is an antagonism between selection pressure against mutants and growth by the merging of mutant domains with an ensuing absorbing state phase transition to an all-mutant front. We find that surface roughening has a marked effect on the critical properties of the absorbing state phase transition. While reference models, which keep the expanding front flat, exhibit directed percolation critical behavior, the exponents of the two-species Eden model strongly deviate from it. In turn, the mutation-selection process induces an increased surface roughness with exponents distinct from that of the classical Eden model