The influence of mesoscale and submesoscale heterogeneity on ocean biogeochemical reactions

The oceanic circulation in the meso to submesoscale regime generates heterogeneity in the concentrations of biogeochemical components over these scales, horizontally between 1 and 100 km. Due to nonlinearities in the biogeochemical reactions, such as phytoplankton primary production and zooplankton grazing, this small-scale heterogeneity can lead to departure from the mean field approximation, whereby plankton reactions are evaluated from mean distributions at coarser scale. Here we explore the magnitude of these eddy reactions and compare their strength to those of the more widely studied eddy transports. We use the term eddy to denote effects arising from scales smaller than approximate to 100 km. This is done using a submesoscale permitting biogeochemical model, representative of the seasonally varying subtropical and subpolar gyres. We found that the eddy reactions associated with primary production and grazing account for 5-30% of productivity and grazing, respectively, depending on location and time of year, and are scale dependent: two thirds are due to heterogeneities at scales 30-100 km and one third to those at scales below 30 km. Moreover, eddy productivities are systematically negative, implying that production tends to be reduced by nonlinear interactions at the mesoscale and smaller. The opposite result is found for eddy grazing, which is generally positive. The contrasting effects result from vertical advection, which negatively correlates phytoplankton and nutrients and positively correlates phytoplankton and zooplankton in the meso to submesoscale range. Moreover, our results highlight the central role played by eddy reactions for ecological aspects and the distribution of organisms and by eddy transport for biogeochemical aspects and nutrient budgets.