Carbon and nitrogen flows during a bloom of the coccolithophore Emiliania huxleyi: Modelling a mesocosm experiment

A dynamic model has been developed to represent biogeochemical variables and processes observed during experimental blooms of the coccolithophore Emiliania huxleyi induced inmesocosms over a period of 23 days. The model describes carbon (C), nitrogen (N), and phosphorus (P) cycling through E. huxleyi and the microbial loop, and computes pH and the partial pressure of carbon dioxide (pCO2) from dissolved inorganic carbon (DIC) and total alkalinity (TA). The main innovations are: 1) the representation of E. huxleyi dynamics using an unbalanced growthmodel in carbon and nitrogen, 2) the gathering of formulations describing typical processes involved in the export of carbon such as primary production, calcification, cellular dissolved organic carbon (DOC) excretion, transparent exopolymer (TEP) formation and viral lyses, and 3) an original and validated representation of the calcification process as a function of the net primary production with a modulation by the intra-cellular N:C ratio mimicking the effect of nutrients limitation on the onset of calcification. It is shown that this new mathematical formulation of calcification provides a better representation of the dynamics of TA, DIC and calcification rates derived from experimental data compared to classicaly used formulations (e.g. function of biomass or of net primary production without anymodulation term). In a first step, the model has been applied to the simulations of present pCO2 conditions. It adequately reproduces the observations for chemical and biological variables and provides an overall view of carbon and nitrogen dynamics. Carbon and nitrogen budgets are derived from the model for the different phases of the bloom, highlighting three distinct phases, reflecting the evolution of the cellular C:N ratio and the interaction between hosts and viruses. During the first phase, inorganic nutrients are massively consumed by E. huxleyi increasing its biomass. Uptakes of carbon and nitrogen are maintained at a constant ratio. The second phase is triggered by the exhaustion of phosphate (PO4 3−). Uptake of carbon and nitrogen being uncoupled, the cellular C:N ratio of E. huxleyi increases. This stimulates the active release of DOC, acting as precursors for TEP. The third phase is characterised by an enhancement of the phytoplankton mortality due to viral lysis. A huge amount of DOC has been accumulated in the mesocosm