In earth system modeling, biogeochemistry strongly affects mass and energy exchanges between ecosystems and the physical climate system (Heinze et al., 2019) . Morphologically, biogeochemistry has three pillars: biology, geophysics, and chemistry. In the context of mathematical modeling, geophysics and chemistry generally have much stronger theoretical foundations than biology (Brutsaert, 2005; Stumm & Morgan, 1996; Vallis, 2006) , even though all three are macroscale responses that emerge from

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... atomic interactions, which in an ideal (but impractical) scenario can be predicted by solving the Schrödinger equation of all atoms together (so that arguably they all are subtopics of physics; Feynman et al., 2011c) . In seeking a better understanding of ecological dynamics, for example, competition and symbiosis, mathematical formulations of the substrate-consumer relationship (e.g., the interactions between many microbes as consumers and their diverse substrates) are essential for theoretical modeling and interpreting empirical experiments, such as phytoplankton population dynamics (Tilman, 1982) , plant-microbial competition of nutrients (Zhu et al., 2017) , and microbial decomposition of organic matter (Tang & Riley, 2013b; Yu