Unbalanced reduction of nutrient loads has created an offshore gradient from phosphorus to nitrogen limitation in the N orth S ea

Measures to reduce eutrophication have often led to a more effective decline of phosphorus (P) than nitrogen (N) concentrations. The resultant changes in riverine nutrient loads can cause an increase in the N : P ratios of coastal waters. During four research cruises along a 450 km transect, we investigated how reductions in nutrient inputs during the past 25 yr have affected nutrient limitation patterns in the North Sea. This revealed a strong offshore gradient of dissolved inorganic N : P ratios in spring, from 375 : 1 nearshore toward 1 : 1 in the central North Sea. This gradient was reflected in high nearshore N : P and C : P ratios of particulate organic matter (mainly phytoplankton), indicative of severe P deficiency of coastal phytoplankton, which may negatively affect higher trophic levels in the food web. Nutrient enrichment bioassays performed on-board showed P and Si limitation of phytoplankton growth nearshore, co-limitation of N and P in a transitional region, and N limitation in the outer-shore waters, confirming the existence of an offshore gradient from P to N limitation. Different species were limited by different nutrients, indicating that further reductions of P loads without concomitant reductions of N loads will suppress colonial Phaeocystis blooms, but will be less effective in diminishing harmful algal blooms by dino- and nanoflagellates. Hence, our results provide evidence that de-eutrophication efforts in northwestern Europe have led to a large imbalance in the N : P stoichiometry of coastal waters of the North Sea, with major consequences for the growth, species composition, and nutritional quality of marine phytoplankton communities.