High bacterivory by the smallest phytoplankton in the North Atlantic Ocean

Planktonic algae, <5 mu m in size are major fixers of inorganic carbon in the ocean(1). They dominate phytoplankton biomass in post-bloom, stratified oceanic temperate waters(2). Traditionally, large and small algae are viewed as having a critical growth dependence on inorganic nutrients, which the latter can better acquire at lower ambient concentrations owing to their higher surface area to volume ratios(3,4). Nonetheless, recent phosphate tracer experiments in the oligotrophic ocean(5) have suggested that small algae obtain inorganic phosphate indirectly, possibly through feeding on bacterioplankton. There have been numerous microscopy- based studies of algae feeding mixotrophically(6,7) in the laboratory(8-10) and field(11-14), as well as mathematical modelling of the ecological importance of mixotrophy(15). However, because of methodological limitations(16) there has not been a direct comparison of obligate heterotrophic and mixotrophic bacterivory. Here we present direct evidence that small algae carry out 40 - 95% of the bacterivory in the euphotic layer of the temperate North Atlantic Ocean in summer. A similar range of 37 - 70% was determined in the surface waters of the tropical North- East Atlantic Ocean, suggesting the global significance of mixotrophy. This finding reveals that even the smallest algae have less dependence on dissolved inorganic nutrients than previously thought, obtaining a quarter of their biomass from bacterivory. This has important implications for how we perceive nutrient acquisition and limitation of carbon- fixing protists as well as control of bacterioplankton in the ocean.