Thermodynamic constraints on the diversity of microbial ecosystems
Most biological processes are driven by non-equilibrium thermodynamics, but despite significant progress in theoretical ecology the constraints this places on ecosystem dynamics has been barely considered. Microbial ecosystems represent a natural place to begin this consideration, as many of the ways they interact, such as metabolite production and cross-feeding, can be described in thermodynamic terms. Previous work considered the impact of thermodynamics such as the rate-yield trade-off on individual species' competitive ability, but restrained from analyzing complex dynamical systems. To address this gap we developed a thermodynamic microbial consumer-resource model with fully reversible reaction kinetics, which allows direct consideration of free-energy dissipation. Using this model, we show that ecosystem diversity increases with supplied free energy, because greater availability of free energy allows for faster ecosystem development. Thus, when species from the initial community begin to go extinct more possible niches have been formed, facilitating increased diversity. Our model also shows that the inclusion of species utilising near-to-equilibrium reactions increases diversity under conditions of low free-energy supply. At low free energy supply thermodynamic interaction types reach comparable strength to the conventional (competition and facilitation) interactions yielding a more nuanced classification of interactions, and emphasising the key role thermodynamics plays in the dynamics. Though our model is valid for all microbial ecosystems where diversification from an initial substrate occurs it is of particular use when the initial substrate is recalcitrant (low-free energy).