Spatial Structure Can Decrease Symbiotic Cooperation

Mutualisms occur when at least two species provide a net fitness benefit to each other. These types of interactions are ubiquitous in nature, with more being discovered regularly. Mutualisms are vital to humankind: Pollinators and soil microbes are critical in agriculture, bacterial microbiomes regulate our health, and domesticated animals provide us with food and companionship. Many hypotheses exist on how mutualisms evolve; however, they are difficult to evaluate without bias, due to the fragile and idiosyncratic systems most often investigated. Instead, we have created an artificial life simulation, Symbulation, which we use to examine mutualism evolution based on (1) the probability of vertical transmission (symbiont being passed to offspring) and (2) the spatial structure of the environment. We found that spatial structure can lead to less mutualism at intermediate vertical transmission rates. We provide evidence that this effect is due to the ability of quasi species to purge parasites, reducing the diversity of available symbionts. Our simulation is easily extended to test many additional hypotheses about the evolution of mutualism and serves as a general model to quantitatively compare how different environments affect the evolution of mutualism.