Size-fractionated dissolved primary production and carbohydrate composition of the coccolithophore Emiliania huxleyi

Extracellular release (ER) by phytoplankton is the major source of fresh dissolved organic carbon (DOC) in marine ecosystems and accompanies primary production during all growth phases. Little is known, so far, on size and composition of released molecules, and to which extent ER occurs passively, by leakage, or actively, by exudation. Here, we report on ER by the widespread and bloom-forming coccolithophore Emiliania huxleyi grown under steady-state conditions in phosphorus-controlled chemostats (N : P = 29, growth rate of mu = 0.2 d(-1)) at present-day and high-CO2 concentrations. C-14 incubations were performed to determine primary production (PP), comprised of particulate ((POC)-C-14) and dissolved organic carbon ((DOC)-C-14). Concentration and composition of particulate combined carbohydrates (pCCHO) and high-molecular-weight (> 1 kDa, HMW) dissolved combined carbohydrates (dCCHO) were determined by ion chromatography. Information on size distribution of ER products was obtained by investigating distinct size classes (< 0.4 mu m ((DOC)-C-14), < 0.45 mu m (HMW-dCCHO), < 1000, < 100 and < 10 kDa) of (DOC)-C-14 and HMW-dCCHO. Our results revealed relatively low ER during steady-state growth, corresponding to similar to 4.5% of primary production, and similar ER rates for all size classes. Acidic sugars had a significant share on freshly produced pCCHO as well as on HMW-dCCHO. While pCCHO and the smallest size fraction (< 10 kDa) of HMW-dCCHO exhibited a similar sugar composition, dominated by high percentage of glucose (74-80 mol %), the composition of HMW-dCCHO size classes > 10 kDa was significantly different, with a higher mol% of arabinose. The mol% of acidic sugars increased and that of glucose decreased with increasing size of HMW-dCCHO. We conclude that larger polysaccharides follow different production and release pathways than smaller molecules, potentially serving distinct ecological and biogeochemical functions.