The spatial and metabolic basis of colony size variation.

Spatial structure impacts microbial growth and interactions, with ecological and evolutionary consequences. It is therefore important to quantitatively understand how spatial proximity affects interactions in different environments. We tested how proximity influences colony size when either Escherichia coli or Salmonella enterica are grown on various carbon sources. The importance of colony location changed with species and carbon source. Spatially explicit, genome-scale metabolic modeling recapitulated observed colony size variation. Competitors that determine territory size, according to Voronoi diagrams, were the most important drivers of variation in colony size. However, the relative importance of different competitors changed through time. Further, the effect of location increased when colonies took up resources quickly relative to the diffusion of limiting resources. These analyses made it apparent that the importance of location was smaller than expected for experiments with S. enterica growing on glucose. The accumulation of toxic byproducts appeared to limit the growth of large colonies and reduced variation in colony size. Our work provides an experimentally and theoretically grounded understanding of how location interacts with metabolism and diffusion to influence microbial interactions.