Title: Contribution of heterotrophic plankton to nitrogen regeneration in the upwelling ecosystem of A Coruna (NW Spain)
Abstract: The contribution of heterotrophic plankton to nitrogen (N) regeneration in the water column, and its significance for the requirements of phytoplankton, were studied at the seasonal scale in the coastal upwelling ecosystem of A Coruna (Galicia, NW Spain). During 1995-1997, monthly measure- ments were taken of hydrographic conditions, dissolved nutrients, and abundance and biomass of microplanktonic heterotrophs (bacteria, flagellates and ciliates), phytoplankton and mesozooplankton (>200 m). Additionally, series of experiments were conducted to quantify N fluxes, including primary production ( 14 C method), phytoplankton uptake of nitrate, ammonium and urea ( 15 N- labelling techniques), microheterotrophic regeneration of ammonium, mesozooplankton grazing (chloro- phyll gut-content method) and excretion of ammonium by mesozooplankton. Two N budgets were built for the average situations of high (>100 mg C m � 2 h � 1 ) and low (<100 mg C m � 2 h � 1 ) primary production. The results revealed that phytoplankton relied strongly on regenerated ammo- nium all year round (33 and 43% of total N uptake in high and low production situations, respectively). This demand for ammonium was closely matched by regeneration rates of micro- plankton (0.14-0.25 mmol N m � 2 h � 1 ), whereas zooplankton contributed on average <10% to N regeneration. Likewise, zooplankton grazing had little direct control on phytoplanktonic biomass. The results obtained indicate that in the A Coruna upwelling system, N biomass of heterotrophic plankton is generally higher than phytoplankton N biomass. The high rates of N regeneration measured also suggest that a large proportion of the organic matter produced after an upwelling pulse is recycled in the water column through the microbial food web.