  Cooperation is ubiquitous in nature, but explaining its existence remains a
central interdisciplinary challenge. Cooperation is most difficult to explain
in the Prisoner's Dilemma game, where cooperators always lose in direct
competition with defectors despite increasing mean fitness. Here we demonstrate
how spatial population expansion, a widespread natural phenomenon, promotes the
evolution of cooperation. We engineer an experimental Prisoner's Dilemma game
in the budding yeast Saccharomyces cerevisiae to show that, despite losing to
defectors in nonexpanding conditions, cooperators increase in frequency in
spatially expanding populations. Fluorescently labeled colonies show genetic
demixing of cooperators and defectors, followed by increase in cooperator
frequency as cooperator sectors overtake neighboring defector sectors. Together
with lattice-based spatial simulations, our results suggest that spatial
population expansion drives the evolution of cooperation by (1) increasing
positive genetic assortment at population frontiers and (2) selecting for
phenotypes maximizing local deme productivity. Spatial expansion thus creates a
selective force whereby cooperator-enriched demes overtake neighboring
defector-enriched demes in a "survival of the fastest". We conclude that colony
growth alone can promote cooperation and prevent defection in microbes. Our
results extend to other species with spatially restricted dispersal undergoing
range expansion, including pathogens, invasive species, and humans.
