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To mitigate coastal eutrophication, a worldwide concern to sustain ecosystem services, actions often focus on nutrient-load reductions but sometimes also aim to limit the abundance of forage fish. Adding to the eutrophication problem, changes in climate and land use lead to increased inputs of coloured dissolved organic matter (darkening). We conducted a mesocosm experiment to study the effects of water darkening on the outcome of eutrophication mitigation actions acting either bottomup or via top-down control. We initiated eutrophication under varying water colour conditions. Then, we performed mitigation actions by stopping nutrient additions, removing forage fish, or both. Stopping nutrient additions was the most effective action in reducing pelagic algal biomass, irrespective of water colour. In contrast, the effect of fish removal varied with water colour, only having an effect in darker waters. We link these varying responses to shifts in nutrient availability and zooplankton community biomass and composition. Our experiment demonstrates that future eutrophication management may need to consider multiple concurrent pressures, including coastal darkening, as they can generate indirect effects caused by trophic interactions.

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.

Coastal eutrophication due to excessive anthropogenic nutrient loading is a major threat worldwide, and especially in estuaries and semi-enclosed waterbodies, like the brackish Baltic Sea. In addition, coastal waters may become darker (coastal darkening) due to increased input of colored compounds from terrestrial run-off and sediment resuspension. Still, the effects of darkening on coastal food web responses to eutrophication are unknown. In a mesocosm experiment with benthic and pelagic habitats, we manipulated nutrient loading, presence of fish and light availability to disentangle bottom-up and top-down control of eutrophication symptoms in ambient and darkened waters. Overall, we found higher pelagic Chlorophyll a concentrations (a proxy of algal biomass) with darkening and with nutrient enrichment in both clear and dark waters. Albeit fish had a strong impact on zooplankton and zoobenthos, they had no cascading effect on algae. We conclude that coastal darkening due to changes in land use and climate change can pose an additional challenge concerning the recovery of coastal waters from eutrophication.

We tested interactive effects of warming (+2°C) and browning on periphyton accrual and pigment composition when grown on a synthetic substrate (plastic strips) in the euphotic zone of 16 experimental ponds. We found that increased colored dissolved organic matter (cDOM) and associated nutrients alone, or in combination with warming, resulted in a substantially enhanced biomass accrual of periphyton, and a comparatively smaller increase in phytoplankton. This illustrates that periphyton is capable of using nutrients associated with cDOM, and by this may affect nutrient availability for phytoplankton. However, warming weakened the positive impact of browning on periphyton accrual, possibly by thermal compensation inferred from altered pigment composition, and/or changes in community composition. Our results illustrate multiple impacts of climate change on algal growth, which could have implications for productivity and consumer resource use, especially in shallow areas in northern lakes.

Benthic gross primary production (GPP) is often the most important part of aquatic food webs in northern lakes, which are gradually warming and receiving increased terrestrial colored dissolved organic carbon loadings due to global change. Yet, measurements of benthic GPP are fairly uncommon, and methods and unit dimensions of benthic GPP are unstandardized and rarely compared. In this study, we measured benthic GPP in 27 headwater lakes from three regions in northern Sweden and analyzed potential constraining drivers of benthic GPPz rates at discrete depths and estimates of benthic GPP averages across the whole lake, as well as across the littoral zone. We also compared in situ measurements of benthic GPP averages across the whole lake with modeled values using the “autotrophic structuring model.” We found that benthic GPPz rates were best explained by, and positively related to, available light (i.e., a function of depth and water color) and temperature. Benthic GPP averages across the whole lake, on the contrary, were best explained by the relative size of the littoral area, which is a measure that combines lake bathymetry and water color. The comparison between in situ measured and modeled estimates of benthic GPP averages across the whole lake revealed that (1) the autotrophic structuring model underestimates GPP at low values and overestimates GPP at high values compared with measured data, and that (2) measured values were related to temperature, which is not included as a variable in the autotrophic structuring model. Considering future predicted changes impacting northern latitude lakes, our results suggest that increased lake water temperatures can to some extent mitigate the negative impacts of reduced light availability from lake browning on benthic GPPz rates. The combined impact of these changes on benthic GPP averages across the whole lake will depend on, and be moderated by, lake bathymetry determining the relative size of the littoral area.

Gross primary production (GPP) by benthic microalgae growing on soft sediments is an important contributor to lake productivity in many lakes world-wide. As benthic microalgae have access to nutrients in the sediment they have been regarded as primarily controlled by light, while the role of CO2 as a limiting factor for benthic GPP in lake ecosystems is largely unknown.

In this study, we experimentally tested for CO2 limitation of benthic GPP by collecting littoral surface sediments, with associated benthic microalgae, from a typical boreal lake. Intact sediment cores were incubated at different depths (light conditions) after addition of dissolved inorganic (bicarbonate) or organic (DOC; glucose) carbon as direct and indirect sources of CO2, respectively.

Benthic microalgal GPP was stimulated by both dissolved inorganic carbon and DOC additions at high, but not at low, light levels.

This study shows that benthic microalgal GPP can be CO2-limited when light is not limiting and suggests that both direct (e.g., via groundwater inflow) and indirect (via mineralisation of DOC) CO2 supply can stimulate benthic GPP.

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.

Rivers transport large amounts of allochthonous organic matter (OM) to the ocean every year, but there are still fundamental gaps in how allochthonous OM is processed in the marine environment. Here, we estimated the relative contribution of allochthonous OM (allochthony) to the biomass of benthic and pelagic consumers in a shallow coastal ecosystem in the northern Baltic Sea. We used deuterium as a tracer of allochthony and assessed both temporal variation (monthly from May to August) and spatial variation (within and outside river plume). We found variability in allochthony in space and time and across species, with overall higher values for zoobenthos (26.2 +/- 20.9%) than for zooplankton (0.8 +/- 0.3%). Zooplankton allochthony was highest in May and very low during the other months, likely as a result of high inputs of allochthonous OM during the spring flood that fueled the pelagic food chain for a short period. In contrast, zoobenthos allochthony was only lower in June and remained high during the other months. Allochthony of zoobenthos was generally higher close to the river mouth than outside of the river plume, whereas it did not vary spatially for zooplankton. Last, zoobenthos allochthony was higher in deeper than in shallower areas, indicating that allochthonous OM might be more important when autochthonous resources are limited. Our results suggest that climate change predictions of increasing inputs of allochthonous OM to coastal ecosystems may affect basal energy sources supporting coastal food webs.

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.

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.