Iron fertilization and the structure of planktonic communities in high nutrient regions of the Southern Ocean
In this review article, plankton community structure observations are analyzed both for artificial iron fertilization experiments and also for experiments dedicated to the study of naturally iron-fertilized systems in the Atlantic, Indian and Pacific sectors of the Southern Ocean in the POOZ (Permanently Open Ocean Zone) and the PFZ (Polar Frontal Zone). Observations made in natural systems are combined with those from artificially perturbed systems, in order to evaluate the seasonal evolution of pelagic communities, taking into account controlling factors related to the life cycles and the ecophysiology of dominant organisms. The analysis considers several types of planktonic communities, including both autotrophs and heterotrophs. These communities are spatially segregated owing to different life strategies. A conceptual general scheme is proposed to account for these observations and their variability, regardless of experiment type. Diatoms can be separated into 2 groups: Group 1 has slightly silicified fast growing cells that are homogeneously distributed in the surface mixed layer, and Group 2 has strongly silicified slowly growing cells within discrete layers. During the growth season, Group 1 diatoms show a typical seasonal succession of dominant species, within time windows of development that are conditioned by physical factors (light and temperature) as well as endogenous specific rhythms (internal clock), and biomass accumulation is controlled by the availability of nutrients. Group 1 diatoms are not directly grazed by mesozooplankton which is fed by protozooplankton, linking the microbial food web to higher trophic levels. Instead, successive dominant species of Group 1 are degraded via bacterial activity at the end of their growth season. Organic detritus fragments feed protozooplankton and mesozooplankton. The effective silicon pump leads to the progressive disappearance of silicic acid in surface waters. In contrast, Group 2 is resistant to grazing due to its strong silicification, and its biomass accumulates continuously but relatively slowly throughout the productive period. Group 2 diatoms are concentrated at or near the seasonal pycnocline and thus benefit from upward nutrient fluxes by diapycnal mixing. The decrease in light and the deep convective mixing in the fall produce both light and nutrient limitation leading to a massive carbon export of Group 2 diatoms, a major annual event of the biological pump. This scheme describes the seasonal evolution of plankton communities in surface waters of the Southern Ocean. The scheme could probably be extended to ecosystems that are characterized by a seasonal bloom under influence of iron or other nutrients.