Understanding microbial cooperation

The field of microbial cooperation has grown enormously over the last decade, leading to improved experimental techniques and a growing awareness of collective behavior in microbes. Unfortunately, many of our theoretical tools and concepts for understanding cooperation fail to take into account the peculiarities of the microbial world, namely strong selection strengths, unique population structure, and non-linear dynamics. Worse yet, common verbal arguments are often far removed from the math involved, leading to confusion and mistakes. Here, we review the general mathematical forms of Price's equation, Hamilton's rule, and multilevel selection as they are applied to microbes and provide some intuition on these otherwise abstract formulas. However, these sometimes overly general equations can lack specificity and predictive power, ultimately forcing us to advocate for more direct modeling techniques.