Planktonic ecosystems provide a key mechanism for the transfer of carbon from the atmosphere to the deep ocean via the so-called "biological pump." Mathematical models of these ecosystems have been used to predict CO2 uptake in surface waters at particular locations, and more recently have been embedded in global climate models. While the equilibrium properties of these models are well studied, less attention has been paid to their response to external perturbations, despite the fact that as a result of the variability of environmental forcing such ecosystems are rarely, if ever, in equilibrium. In this study, linear theory is used to determine the structure of perturbations to state variables of an ecosystem model describing summertime conditions at Ocean Station P (50°N 145°W) that maximize either instantaneous or integrated export flux. As a result of the presence of both direct and indirect pathways to export in this model, these perturbations involve the dynamics of the entire ecosystem. For all "optimal" perturbations considered, it is found that the flux to higher trophic levels is the primary contributor to export flux, followed by sinking detritus. In contrast, the contribution of aggregation is negligible. In addition, small phytoplankton contribute significantly (comparable to large phytoplankton) to the export flux through indirect pathways, primarily through the microzooplankton, even following a bloom in only large phytoplankton. While the details of these results may be specific to the particular model under consideration, the optimal perturbation framework is general and can be used to probe the dynamics of any mechanistic ecosystem model.
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