Carbon limitation of lake productivity.

Phytoplankton productivity in lakes controls the rate of synthesis of organic matter that drives energy flow through the food webs and regulates the transparency and oxygen conditions in the water. Limitation of phytoplankton productivity and biomass by nutrients and light availability is an established paradigm for lake ecosystems, whereas invasion of atmospheric CO2 has been assumed to cover the high demands of dissolved inorganic carbon (DIC) during intense organic productivity. We challenge this paradigm, and show up to a 5-fold stimulation of phytoplankton productivity and biomass in outdoor mesocosms enriched with DIC, compared to mesocosms with lower DIC concentrations. High DIC supported phytoplankton productivity by direct algal uptake of bicarbonate, through the release of CO2 coupled to calcification and by inducing high pH that greatly enhances atmospheric CO2 invasion. Comparisons of 204 natural Danish lakes supported mesocosm experiments showing higher phytoplankton biomass and pH levels in hard water than soft water lakes for the same nutrient and light availabilities. The most productive lakes are nutrient-rich, hard water lakes that attain surface pHs of 10-11 and chemically enhance atmospheric CO2 uptake 10-15-fold. Our results will help understand natural variations of lake productivity along gradients in nutrients, DIC and pH.