Impact of Dynamic Phytoplankton Stoichiometry on Global Scale Patterns of Nutrient Limitation, Nitrogen Fixation, and Carbon Export

Abstract

Phytoplankton stoichiometry modulates the interaction between carbon, nitrogen and phosphorus cycles. Environmentally driven variations in phytoplankton C:N:P can alter biogeochemical cycling compared to expectations under fixed ratios. In fact, the assumption of fixed C:N:P has been linked to Earth System Model (ESM) biases and potential misrepresentation of responses to future change. Here we integrate key elements of the Adaptive Trait Optimization Model (ATOM) for phytoplankton stoichiometry with the Carbon, Ocean Biogeochemistry and Lower Trophics (COBALT) ocean biogeochemical model. Within a series of global ocean-ice-ecosystem retrospective simulations, ATOM-COBALT reproduced observations of phytoplankton N:P, and compared to static ratios, exhibited reduced phytoplankton P-limitation, enhanced N-fixation, and increased low-latitude export, improving consistency with observations and highlighting the biogeochemical implications of dynamic N:P. We applied ATOM-COBALT to explore the impacts of different physiological mechanisms hypothesized to underlie N:P variation, finding that two mechanisms together drove the observed patterns: proportionality of P-rich ribosomes in phytoplankton cells to growth rates and reductions in P-storage during scarcity. A third mechanism which linked temperature with phytoplankton biomass allocations to non-ribosomal proteins, led only to relatively modest impacts because this mechanism decreased the temperature dependence of phytoplankton growth rates, compensating for changes in N:P. We find that there are quantitative response differences that associate distinctive biogeochemical footprints with each mechanism, which are most apparent in highly productive low-latitude regions. These results suggest that variable phytoplankton N:P makes phytoplankton productivity and export resilient to environmental changes, and support further research on the physiological and environmental drivers of phytoplankton stoichiometry and biogeochemical role.