Global change has increased inorganic nitrogen (N) and dissolved organic carbon (DOC; i.e., "browning") inputs to northern hemisphere boreal lakes. However, we do not know how phytoplankton in nutrient poor lake ecosystems of different DOC concentration respond to increased N availability. Here, we monitored changes in phytoplankton production, biomass and community composition in response to whole lake inorganic N fertilization in six boreal unproductive Swedish lakes divided into three lake pairs (control, N enriched) at three DOC levels (low, medium, high), with one reference year (2011) and 2 impact yr (2012, 2013). We found that phytoplankton biomass and production decreased with DOC concentration before N fertilization. Further, phytoplankton community composition also differed with respect to DOC, with a dominance of non-flagellated autotrophs at low DOC towards an increasing dominance of flagellated autotrophs with increased lake DOC concentration. The N fertilization increased phytoplankton biomass and production in all lakes, but did not affect phytoplankton community composition. However, the net response in biomass and production to N fertilization declined with increasing DOC, implying that the lake DOC concentration is critical in order to infer phytoplankton responses to N fertilization, and that the system switches from being primarily nutrient limited to becoming increasingly light limited with increased DOC concentration. In conclusion, our results show that browning will reduce phytoplankton production and biomass and influence phytoplankton community composition, whereas increased inorganic N loadings from deposition, forestry or other land use will primarily enhance phytoplankton biomass and production. Together, any change in the landscape that enhances inorganic N availability will increase phytoplankton production and biomass, but the positive effects of N will be much weaker or even neutralized in browner lakes as caused by light limitation.

The aim of this study was to predict the combined effects of enhanced nitrogen (N) deposition and warming on phytoplankton development in high latitude and mountain lakes. Consequently, we assessed, in a series of enclosure experiments, how lake water nutrient stoichiometry and phytoplankton nutrient limitation varied over the growing season in 11 lakes situated along an altitudinal/climate gradient with low N-deposition (<1 kg N ha(-1)  yr(-1) ) in northern subarctic Sweden. Short-term bioassay experiments with N- and P-additions revealed that phytoplankton in high-alpine lakes were more prone to P-limitation, and with decreasing altitude became increasingly N- and NP-colimited. Nutrient limitation was additionally most obvious in midsummer. There was also a strong positive correlation between phytoplankton growth and water temperature in the bioassays. Although excess nutrients were available in spring and autumn, on these occasions growth was likely constrained by low water temperatures. These results imply that enhanced N-deposition over the Swedish mountain areas will, with the exception of high-alpine lakes, enhance biomass and drive phytoplankton from N- to P-limitation. However, if not accompanied by warming, N-input from deposition will stimulate limited phytoplankton growth due to low water temperatures during large parts of the growing season. Direct effects of warming, allowing increased metabolic rates and an extension of the growing season, seem equally crucial to synergistically enhance phytoplankton development in these lakes.

Anthropogenic activities are increasing inorganic nitrogen (N) loadings to unproductive boreal lakes. In many of these lakes phytoplankton are N limited, consequently N fertilization may affect ecosystem productivity and consumer resource use. Here, we conducted whole lake inorganic N fertilization experiments with six small N limited unproductive boreal lakes (three control and three N enriched) in an area receiving low N deposition with one reference and two impact years. Our aim was to assess the effects of N fertilization on pelagic biomass production and consumer resource use. We found that phytoplankton primary production (PP) and biomass, and the PP: bacterioplankton production ratio increased after fertilization. As expected, the relative contribution of phytoplankton derived resources (autochthony) that supported the crustacean zooplankton community increased. Yet, the response in the consumer community was modest with autochthony only increasing in one of the three major zooplankton groups and with no effect on zooplankton biomass. In conclusion, our findings imply that newly available phytoplankton energy derived from N fertilization was not efficiently transferred up to zooplankton, indicating a mismatch between producer energy supply and consumer energy use with potential accumulation of phytoplankton biomass as the result.

Additions of labile organic carbon (C) enhanced bacterial production (BP) and were associated with increases in crustacean zooplankton and planktivorous fish biomasses. This was shown in a mesocosm experiment where we traced the contribution of BP to zooplankton and fish using stable isotopes and labile glucose-C as a biomarker. BP increased with glucose-C addition, and all zooplankton and fish incorporated some glucose-C. However, the effect of labile-C addition on zooplankton was taxa-dependant, as although cladocerans incorporated the most labile-C, increased BP did not affect cladoceran biomass. Instead, calanoid copepod biomass increased with glucose addition. This suggests that the ability to selectively graze on high quality food, such as bacterial grazing protists capable of trophic upgrading, had a stronger positive effect on calanoid copepods biomass than unselective grazing on bacteria and protists had on cladoceran biomass. Higher BP was associated with increased survival and population growth of young-of-the-year perch (Perca fluviatilis) when stocked at high densities, which suggested that BP had a density-dependant positive effect on fish growth.Résumé : Les additions de carbone (C) organique labile augmentent la production bactérienne (PB) et sont associées à des accroissements des biomasses des crustacés zooplanctoniques et des poissons planctonophages. Cela est bien démontré dans une expérience en microcosme dans laquelle nous avons pu suivre la contribution de la BP au zooplancton et aux poissons à l'aide d'isotopes stables et du C-glucose comme biomarqueur. La BP augmente avec l'addition de C-glucose et tout le zooplancton et les poissons incorporent du C-glucose. Cependant, les effets de l'addition de C labile sur le zooplancton dé-pend des taxons, puisque même si les cladocères incorporent le plus le C labile, l'augmentation de la BP n'affecte pas la biomasse des cladocères. En revanche, la biomasse des copépodes calanoïdes augmente avec l'addition de glucose. Cela laisse penser que la capacité de brouter de façon sélective des aliments de haute qualité, tels que des protistes qui se nourrissent de bactéries et sont capables d'un surclassement trophique, a un effet positif plus fort sur les copépodes calanoïdes, que le broutage non sélectif de bactéries et de protistes peut avoir sur la biomasse des cladocères. La BP plus élevée est associée à une survie et une croissance de population plus fortes des jeunes perches (Perca fluviatilis) de l'année lorsqu'elles sont empoissonnées à forte densité, ce qui laisse croire que la BP a un effet positif dépendant de la densité sur la croissance des poissons.[Traduit par la Rédaction]