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Globally, inland waters emit over 2 Pg of carbon per year as carbon dioxide, of which the majority originates from streams and rivers. Despite the global significance of fluvial carbon dioxide emissions, little is known about their diel dynamics. Here we present a large-scale assessment of day- and night-time carbon dioxide fluxes at the water-air interface across 34 European streams. We directly measured fluxes four times between October 2016 and July 2017 using drifting chambers. Median fluxes are 1.4 and 2.1mmolm(-2) h(-1) at midday and midnight, respectively, with night fluxes exceeding those during the day by 39%. We attribute diel carbon dioxide flux variability mainly to changes in the water partial pressure of carbon dioxide. However, no consistent drivers could be identified across sites. Our findings highlight widespread day-night changes in fluvial carbon dioxide fluxes and suggest that the time of day greatly influences measured carbon dioxide fluxes across European streams. Diel patterns can greatly impact total stream carbon dioxide emissions, with 39% greater carbon dioxide flux during the night-time relative to the day-time, according to a study of 34 streams across Europe.

Forest harvest in the boreal zone can increase the input of terrestrial materials such as dissolved organic carbon (DOC) and nitrate (NO3-) into nearby aquatic ecosystems, with potential effects on phytoplankton growth through enhanced nutrient (i.e., positive) or reduced light availability (i.e., negative), which may affect ecosystem productivity and consumer resource use. Here, we conducted forest clear-cutting experiments in the catchments of four small, humic, and nitrogen-limited unproductive boreal lakes (two controls and two clear-cut, 18% and 44% of area cut) with one reference and two impact years. Our aim was to assess the effects of forest clear-cutting on pelagic biomass production and consumer resource use. We found that pelagic biomass production did not change after two years of forest clear-cutting: Pelagic primary and bacterial production (PP, BP), PP:BP ratio, chl a, and seston carbon (seston C) were unaffected by clear-cutting; neither did tree harvest affect seston stoichiometry (i.e., N:phosphorus [P], C:P) nor induce changes in zooplankton resource use, biomass, or community composition. In conclusion, our findings suggest that pelagic food webs of humic lakes (DOC > 15 mg/L) might be resilient to a moderate form of forest clear-cutting, at least two years after tree removal, before mechanical site preparation (e.g., mounding, plowing) and when leaving buffer strips along lakes and incoming streams. Thus, pelagic food web responses to forest clear-cutting might not be universal, but could depend on factors such as the time scale, share of catchment logged, and the forest practices involved, including the application of buffer strips and site preparation.

Dietary uptake is a key step in conveying both toxic mercury (Hg; particularly as highly bioavailable methylmercury, MeHg) and essential dietary biochemicals, such as polyunsaturated fatty acids (PUFA), across trophic levels within aquatic food webs. Using stable isotopes and fatty acids we evaluated the role of food sources in size-fractioned plankton and littoral macroinvertebrates for the bioaccumulation of total Hg and MeHg in six oligotrophic and one mesotrophic Swedish lakes with differing concentrations of dissolved organic carbon (DOC). We found that the consumption of both algal and terrestrial diets (assessed by PUFA and long-chain saturated fatty acids, respectively) predicted >66% of the Hg concentration variability in meso- (100-500 mu m) and macrozooplankton (>500 mu m) in oligotrophic lakes. In the mesotrophic lake, total Hg bioaccumulation in higher trophic level biota, carnivorous macroinvertebrates was also significantly related to terrestrial diet sources (R-2 = 0.65, p < 0.01). However, lake pH and DOC correlated to total Hg bioaccumulation and bioconcentration across all lakes, suggesting the consumption of different diet sources is mediated by the influence of lake characteristics. This field study reveals that using dietary biomarkers (stable isotopes and fatty acids) together with the physico-chemical lake parameters pH and nutrients together improve our ability to predict Hg bioaccumulation in aquatic food webs. Fatty acids used as dietary biomarkers provide correlative evidence of specific diet source retention in consumers and their effect on Hg bioaccumulation, while pH and nutrients are the underlying physico-chemical lake parameters controlling differences in Hg bioaccumulation between lakes. 

Large-scale studies are needed to identify the drivers of total mercury (THg) and monomethyl-mercury (MeHg) concentrations in aquatic ecosystems. Studies attempting to link dissolved organic matter (DOM) to levels of THg or MeHg are few and geographically constrained. Additionally, stream and river systems have been understudied as compared to lakes. Hence, the aim of this study was to examine the influence of DOM concentration and composition, morphological descriptors, land uses and water chemistry on THg and MeHg concentrations and the percentage of THg as MeHg (%MeHg) in 29 streams across Europe spanning from 41°N to 64°N. THg concentrations (0.06–2.78 ng L⁻¹) were highest in streams characterized by DOM with a high terrestrial soil signature and low nutrient content. MeHg concentrations (7.8–159 pg L⁻¹) varied non-systematically across systems. Relationships between DOM bulk characteristics and THg and MeHg suggest that while soil derived DOM inputs control THg concentrations, autochthonous DOM (aquatically produced) and the availability of electron acceptors for Hg methylating microorganisms (e.g. sulfate) drive %MeHg and potentially MeHg concentration. Overall, these results highlight the large spatial variability in THg and MeHg concentrations at the European scale, and underscore the importance of DOM composition on mercury cycling in fluvial systems.

Global warming and land use scenarios suggest increased 21^st century nitrogen (N) inputs to aquatic systems. Nitrogen affects in-lake processing and, potentially, atmospheric exchange of greenhouse gases, probably being most relevant in unproductive systems. Here, we test for the first time the effect of a whole-lake experimental increase (threefold) in external nitrate loads on the atmospheric exchange of carbon dioxide (CO₂), methane (CH₄), and nitrous oxide (N₂O) from N-limited unproductive boreal lakes. Nitrate enrichment effects were assessed within a paired Before/After-Control/Impact framework based on 2-hourly to biweekly surface-water sampling of dissolved gas concentrations, and monthly whole-lake inventory surveys, carried out over 4 yrs in six lakes. Nitrate enrichment did not affect gas exchange during summer stratification and whole-lake gas inventories during summer and winter stratification. This finding specifically emphasizes the modest role of internal carbon fixation for the CO₂ dynamics of unproductive boreal lakes. A global synthesis of 52 published studies revealed a wide range of nutrient fertilization effects, both in systems similar to our experimental lakes, and other more productive systems. Effects depended mainly on the spatiotemporal scale of the study and became more pronounced when N enrichment was combined with phosphorous. Conclusively, although short-term and habitat-specific effects can occur, changes in N supply have only weak whole-ecosystem effects on greenhouse gas emissions from unproductive boreal lakes.

Anthropogenic activities are increasing inorganic nitrogen (N) loadings to lakes in the northern hemisphere. In many boreal lakes phytoplankton are N limited, wherefore enhanced N input may affect the productivity of pelagic food webs. Simultaneously, global change causes increased inflows of terrestrial dissolved organic carbon (DOC) to boreal lakes. Between clear and humic lakes, whole lake primary and consumer production naturally differs. However, research is inconclusive as to what controls pelagic production in these lakes. Further, it is unclear how DOC affects the response of the pelagic food web to enhanced inorganic N availability. The overarching goal of this thesis was to study the effects of inorganic N and organic C for pelagic food webs in boreal lakes. In the thesis, I first identified the main drivers of pelagic production during summer in eight non-manipulated Swedish boreal lakes with naturally low or high DOC. Then I investigated how increased N availability affects the pelagic food chain, and how the response differs with DOC. Therefore, whole lake inorganic N fertilization experiments were conducted in six Swedish boreal lakes across a DOC gradient (low, medium, high) divided into three lake pairs (control, N enriched) with one reference and two impact years. In each lake, I also investigated the response of zooplankton growth using in situ mesocosm experiments excluding planktivores. I found that humic boreal lakes had lower phytoplankton production and biomass than clear water lakes. Further, phytoplankton community composition and food quality differed with DOC. However, high DOC did not reduce pelagic energy mobilization or zooplankton biomass, but promoted a higher dominance of cladoceran relative to copepod species. N addition clearly enhanced phytoplankton biomass and production in the experimental lakes. However, this stimulating N effect decreased with DOC as caused by light limitation. Further, the newly available phytoplankton energy derived from N addition was not efficiently transferred to zooplankton, which indicates a mismatch between producer energy supply and consumer energy use. Indeed, the mesocosm experiment revealed that decreased food quality of phytoplankton in response to N addition resulted in reduced food web performance, especially in clearer lakes. In humic lakes, zooplankton production and food web efficiency were clearly more resilient to N addition. In summary, my thesis suggests that any change in the landscape that enhances inorganic N availability will especially affect pelagic food webs in clear water lakes. In contrast, brownification will result in more lakes being resilient to eutrophication caused by enhanced N deposition.

Global environmental change has altered the nitrogen (N) cycle and enhanced terrestrial dissolved organic carbon (DOC) loadings to northern boreal lakes. However, it is still unclear how enhanced N availability affects pelagic food web efficiency (FWE) and crustacean zooplankton growth in N limited boreal lakes. Here, we performed in situ mesocosm experiments in six unproductive boreal Swedish lakes, paired across a DOC gradient, with one lake in each pair fertilized with N (2011: reference year; 2012, 2013: impact years). We assessed how zooplankton growth and FWE were affected by changes in pelagic energy mobilization (PEM), food chain length (phytoplankton versus bacterial production based food chain, i.e. PP:BP), and food quality (seston stoichiometry) in response to N fertilization. Although PP, PEM and PP:BP increased in low and medium DOC lakes after N fertilization, consumer growth and FWE were reduced, especially at low DOC-potentially due to reduced phytoplankton food quality [increased C: phosphorus (P); N:P]. At high DOC, N fertilization caused modest increases in PP and PEM, with marginal changes in PP:BP and phytoplankton food quality, which, combined, led to a slight increase in zooplankton growth and FWE. Consequently, at low DOC (<12 mg L-1), increased N availability lowers FWE due to mismatches in food quality demand and supply, whereas at high DOC this mismatch does not occur, and zooplankton production and FWE may increase. We conclude that the lake DOC level is critical for predicting the effects of enhanced inorganic N availability on pelagic productivity in boreal 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.

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 two impact years (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.

Climate change scenarios predict intensified terrestrial storm runoff, providing coastal ecosystems with large nutrient pulses and increased turbidity, with unknown consequences for the phytoplankton community. We conducted a 12-day mesocosm experiment in the Mediterranean Thau Lagoon (France), adding soil (simulated runoff) and fish (different food webs) in a 2 x 2 full factorial design and monitored phytoplankton composition, shade adaptation and stoichiometry. Diatoms (Chaetoceros) increased fourfold immediately after soil addition, prymnesiophytes and dinoflagellates peaked after six- and 12 days, respectively. Soil induced no phytoplanlcton shade adaptation. Fish reduced the positive soil effect on dinoflagellates (Scripsiella, Glenodinium), and diatom abundance in general. Phytoplankton community composition drove seston stoichiometry. In conclusion, pulsed terrestrial runoff can cause rapid, low quality (high carbon: nutrient) diatom blooms. However, bloom duration may be short and reduced in magnitude by fish. Thus, climate change may shift shallow coastal ecosystems towards famine or feast dynamics.