Browning affects pelagic productivity in northern lakes by surface water warming and carbon fertilization.

Global change impacts important environmental drivers for pelagic gross primary production (GPP) in northern lakes, such as temperature, light, nutrient, and inorganic carbon availability. Separate and/or synergistic impacts of these environmental drivers on pelagic GPP remain largely unresolved. Here, we assess key drivers of pelagic GPP by combining detailed depth profiles of summer pelagic GPP with environmental and climatic data across 45 small and shallow lakes across northern Sweden (20 boreal, 6 subarctic, and 19 arctic lakes). We found that across lakes summer pelagic GPP was strongest associated with lake water temperatures, lake carbon dioxide (CO2 ) concentrations impacted by lake water pH, and further moderated by dissolved organic carbon (DOC) concentrations influencing light and nutrient conditions. We further used this dataset to assess the extent of additional DOC-induced warming of epilimnia (here named internal warming), which was especially pronounced in shallow lakes (decreasing 0.96°C for every decreasing m in average lake depth) and increased with higher concentrations of DOC. Additionally, the total pools and relative proportion of dissolved inorganic carbon and DOC, further influenced pelagic GPP with drivers differing slightly among the boreal, subarctic and Arctic biomes. Our study provides novel insights in that global change affects pelagic GPP in northern lakes not only by modifying the organic carbon cycle and light and nutrient conditions, but also through modifications of inorganic carbon supply and temperature. Considering the large-scale impacts and similarities of global warming, browning and recovery from acidification of lakes at higher latitudes throughout the northern hemisphere, these changes are likely to operate on a global scale.

Landscape determinants of pelagic and benthic primary production in northern lakes.

Global change affects gross primary production (GPP) in benthic and pelagic habitats of northern lakes by influencing catchment characteristics and lake water biogeochemistry. However, how changes in key environmental drivers manifest and impact total (i.e., benthic + pelagic) GPP and the partitioning of total GPP between habitats represented by the benthic share (autotrophic structuring) is unclear. Using a dataset from 26 shallow lakes located across Arctic, subarctic, and boreal northern Sweden, we investigate how catchment properties (air temperature, land cover, hydrology) affect lake physico-chemistry and patterns of total GPP and autotrophic structuring. We find that total GPP was mostly light limited, due to high dissolved organic carbon (DOC) concentrations originating from catchment soils with coniferous vegetation and wetlands, which is further promoted by high catchment runoff. In contrast, autotrophic structuring related mostly to the relative size of the benthic habitat, and was potentially modified by CO2 fertilization in the subarctic, resulting in significantly higher total GPP relative to the other biomes. Across Arctic and subarctic sites, DIC and CO2 were unrelated to DOC, indicating that external inputs of inorganic carbon can influence lake productivity patterns independent of terrestrial DOC supply. By comparison, DOC and CO2 were correlated across boreal lakes, suggesting that DOC mineralization acts as an important CO2 source for these sites. Our results underline that GPP as a resource is regulated by landscape properties, and is sensitive to large-scale global changes (warming, hydrological intensification, recovery of acidification) that promote changes in catchment characteristics and aquatic physico-chemistry. Our findings aid in predicting global change impacts on autotrophic structuring, and thus community structure and resource use of aquatic consumers in general. Given the similarities of global changes across the Northern hemisphere, our findings are likely relevant for northern lakes globally.

Author Correction: Carbon dioxide stimulates lake primary production.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Carbon dioxide stimulates lake primary production.

Gross primary production (GPP) is a fundamental ecosystem process that sequesters carbon dioxide (CO2) and forms the resource base for higher trophic levels. Still, the relative contribution of different controls on GPP at the whole-ecosystem scale is far from resolved. Here we show, by manipulating CO2 concentrations in large-scale experimental pond ecosystems, that CO2 availability is a key driver of whole-ecosystem GPP. This result suggests we need to reformulate past conceptual models describing controls of lake ecosystem productivity and include our findings when developing models used to predict future lake ecosystem responses to environmental change.

Global change-driven effects on dissolved organic matter composition: Implications for food webs of northern lakes.

Northern ecosystems are experiencing some of the most dramatic impacts of global change on Earth. Rising temperatures, hydrological intensification, changes in atmospheric acid deposition and associated acidification recovery, and changes in vegetative cover are resulting in fundamental changes in terrestrial-aquatic biogeochemical linkages. The effects of global change are readily observed in alterations in the supply of dissolved organic matter (DOM)-the messenger between terrestrial and lake ecosystems-with potentially profound effects on the structure and function of lakes. Northern terrestrial ecosystems contain substantial stores of organic matter and filter or funnel DOM, affecting the timing and magnitude of DOM delivery to surface waters. This terrestrial DOM is processed in streams, rivers, and lakes, ultimately shifting its composition, stoichiometry, and bioavailability. Here, we explore the potential consequences of these global change-driven effects for lake food webs at northern latitudes. Notably, we provide evidence that increased allochthonous DOM supply to lakes is overwhelming increased autochthonous DOM supply that potentially results from earlier ice-out and a longer growing season. Furthermore, we assess the potential implications of this shift for the nutritional quality of autotrophs in terms of their stoichiometry, fatty acid composition, toxin production, and methylmercury concentration, and therefore, contaminant transfer through the food web. We conclude that global change in northern regions leads not only to reduced primary productivity but also to nutritionally poorer lake food webs, with discernible consequences for the trophic web to fish and humans.

Importance of coastal primary production in the northern Baltic Sea.

In this study, we measured depth-dependent benthic microalgal primary production in a Bothnian Bay estuary to estimate the benthic contribution to total primary production. In addition, we compiled data on benthic microalgal primary production in the entire Baltic Sea. In the estuary, the benthic habitat contributed 17 % to the total annual primary production, and when upscaling our data to the entire Bothnian Bay, the corresponding value was 31 %. This estimated benthic share (31 %) is three times higher compared to past estimates of 10 %. The main reason for this discrepancy is the lack of data regarding benthic primary production in the northern Baltic Sea, but also that past studies overestimated the importance of pelagic primary production by not correcting for system-specific bathymetric variation. Our study thus highlights the importance of benthic communities for the northern Baltic Sea ecosystem in general and for future management strategies and ecosystem studies in particular.

Viruses in subarctic lakes and their impact on benthic and pelagic bacteria.

Virus-bacterium interactions were investigated in the pelagic and benthic habitats in a set of lakes along an altitudinal gradient in the subarctic northern Sweden. Viral and bacterial abundances showed a significant variation between the lakes, with the highest benthic microbial abundances recorded in a high-altitude lake [993 m above sea level (a.s.l.)], whereas the highest pelagic microbial abundances were found in a low-altitude lake (270 m a.s.l.). In the pelagic habitat, there was also a distinct difference in microbial abundances between the summer-autumn and the winter sampling occasion. A positive relationship was noted between viruses and bacteria in both the pelagic and the benthic habitats. Visibly virus-infected bacterial cells were uncommon in the pelagic habitat and undetectable in the benthos. Both lytic and lysogenic pelagic viral production rates were undetectable or low; thus, a possible explanation for the relative high viral abundances found in the water column could be an allochthonous input of viruses or release of sediment-derived viruses. Overall, our results provide novel information about the relevance of viruses in the subarctic region and indicate that viruses play only a minor role in the nutrient and carbon cycling in the microbial communities of subarctic lakes.

Whole-lake estimates of carbon flux through algae and bacteria in benthic and pelagic habitats of clear-water lakes.

This study quantified new biomass production of algae and bacteria in both benthic and pelagic habitats of clear-water lakes to contrast how carbon from the atmosphere and terrestrial sources regulates whole-lake metabolism. We studied four small unproductive lakes in subarctic northern Sweden during one summer season. The production of new biomass in both benthic and pelagic habitats was calculated as the sum of autotrophic production by algae and heterotrophic production by bacteria using allochthonous organic carbon (OC). Whole-lake production of new biomass was dominated by the benthic habitat (86% +/- 4% [mean +/- SD]) and by primary production (77% +/- 9%). Still, heterotrophic bacteria fueled by allochthonous OC constituted a significant portion of the new biomass production in both benthic (19% +/- 11%) and pelagic habitats (51% +/- 24%). In addition, overall net production (primary production minus respiration) was close to zero in the benthic habitats but highly negative (-163 +/- 81 mg C x m(-2) x d(-1)) in pelagic regions of all lakes. We conclude (1) that allochthonous OC supported a significant part of total production of new biomass in both pelagic and benthic habitats, (2) that benthic habitats dominated the whole-lake production of new biomass, and (3) that respiration and net CO2 production dominated the carbon flux of the pelagic habitats and biomass production dominated the benthic carbon flux. Taken together, these findings suggest that previous investigations have greatly underestimated the productivity of clear-water lakes when benthic autotrophic production and metabolism of allochthonous OC have not been measured.

Light limitation of nutrient-poor lake ecosystems.

Productivity denotes the rate of biomass synthesis in ecosystems and is a fundamental characteristic that frames ecosystem function and management. Limitation of productivity by nutrient availability is an established paradigm for lake ecosystems. Here, we assess the relevance of this paradigm for a majority of the world's small, nutrient-poor lakes, with different concentrations of coloured organic matter. By comparing small unproductive lakes along a water colour gradient, we show that coloured terrestrial organic matter controls the key process for new biomass synthesis (the benthic primary production) through its effects on light attenuation. We also show that this translates into effects on production and biomass of higher trophic levels (benthic invertebrates and fish). These results are inconsistent with the idea that nutrient supply primarily controls lake productivity, and we propose that a large share of the world's unproductive lakes, within natural variations of organic carbon and nutrient input, are limited by light and not by nutrients. We anticipate that our result will have implications for understanding lake ecosystem function and responses to environmental change. Catchment export of coloured organic matter is sensitive to short-term natural variability and long-term, large-scale changes, driven by climate and different anthropogenic influences. Consequently, changes in terrestrial carbon cycling will have pronounced effects on most lake ecosystems by mediating changes in light climate and productivity of lakes.