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Land-to-water hydrological connections represent a key regulatory mechanism of carbon transport, controlling carbon dioxide (CO2) emissions from lakes; however, as of yet, there is no assessment of its role at a pan-Arctic scale across large climatic and topographical gradients. We hypothesized that hydrologically well-connected lakes in wetter regions are CO2 sources fueled by stronger lateral fluxes of external carbon relative to drier regions. However, based on data from >200 Arctic lakes, we found that lakes in drier regions have higher and more variable annual CO2 emissions (37.06.2146.0 gC m-2 yr-1, medianQ1Q3) compared to lakes in wetter regions (8.01.717.3 gC m-2 yr-1), with both the lowest and the highest fluxes recorded among dryland lakes. We hypothesize that with increasing wetness, the relative proportion of fluvial emissions increases, whereas in drier landscapes where lakes often have limited stream export, carbon inputs can be retained and more efficiently emitted from lakes.

Nitrogen (N) deposition and dissolved organic carbon (DOC) levels in northern lakes are shifting due to climate change and atmospheric deposition declines, altering the availability of light and nutrients in these ecosystems. Yet their impacts on the biomass, stoichiometry, and the structure of planktonic food chains remain uncertain. We therefore investigated zooplankton-to-seston biomass ratios (Z:S in C, N, and P) across 34 Swedish lakes with varying N deposition, DOC concentration, and fish predation control. Mean Z:S values were 2.9% for C, 7.5% for N, and 7.7% for P, with substantial regional variation. Z:S ratios were higher in lakes with lower atmospheric N deposition, improved seston quality, and greater calanoid copepod dominance in zooplankton. The strong link between zooplankton stoichiometry and community composition underscores the role of calanoids in regulating nutrient dynamics in northern lakes. Fish predation reduced zooplankton biomass but did not significantly alter Z:S ratios or zooplankton community composition. Meanwhile, increasing DOC dampened the higher Z:S in low N deposition lakes by reducing calanoid dominance and promoting more uniform zooplankton assemblages. Our findings suggest that lake browning counteracts the expected increase in Z:S ratios associated with recovery from atmospheric N deposition, potentially altering nutrient transfer in lake food webs.

Climate warming is increasing thermal stratification depth, strength, and duration in lakes, leading to more frequent hypolimnetic oxygen depletion. Most research has focused on eutrophic temperate lakes, which differ significantly from boreal lakes that dominate Earth’s landscape. However, assessing the impact of hypoxia, confounded by browning, warming, and altered stratification, on biogeochemistry and ecological processes in boreal lakes is particularly challenging. Here, we test how oxygenating a hypoxic hypolimnion affects water chemistry, bacterial and primary production (BP and PP), and detritus degradation in a shallow humic boreal lake divided into two basins in an experimental four-year Before-After Control-Impact (BACI) design. After two control years, we oxygenated the hypolimnion of one basin during two stratified periods without disturbing the seasonal development of the thermocline. Hypolimnetic oxygen concentrations moderately impacted lake biogeochemistry. Reoxygenation altered nitrification pathways (increased NO3−) of the hypolimnion, and slightly decreased epilimnion and lake BP (− 6.1% of annual average) and green tea degradation (− 6.0%), whereas Rooibos degradation slightly increased (7.3%). Other water chemistry parameters remained within natural variation. We compared our BACI approach, which separates natural variation, to the simpler Before vs After approach, which does not. We find that studies not accounting for seasonal and among-year variability may overestimate the effects of oxygenation on hypolimnion biogeochemistry, as much of the observed impact is due to natural climate variation. Climate warming and altered stratification patterns are therefore likely to impact boreal lake algal and bacterial production and degradation more than hypolimnion hypoxia during the stratified period.

Widespread increases in lake browning, which affects primary production, have been observed in northern lakes. While lake browning is attributed to increases in terrestrially derived total organic carbon (TOC) and total iron (Fe), Fe does not consistently correlate with increasing TOC over time. This temporal mismatch between TOC and Fe indicates that we still do not fully understand the causes of lake browning, especially in the context of gradually changing climatic conditions. In this study, we utilized Fennoscandian 30-year (1990–2020) time series data for 102 lakes to describe possible reasons for the temporal decoupling between TOC and Fe. Using Bayesian mixed-effects models and wavelet coherence analysis, we found evidence for differential responses of TOC and Fe concentrations to changes in precipitation, temperature, and sulfur deposition. While TOC appeared more sensitive to the effects of precipitation, temperature and sulfur deposition in individual lakes, Fe concentrations were impacted by complex interactions among these environmental variables. Although TOC and Fe increased in most lakes in response to increased temperature and precipitation, 41% of the lakes—typically with larger catchment-to-lake area ratios and shorter water residence times—exhibited a declining trend in Fe. This analysis encompasses lakes of both significant and non-significant changes over time. This decline in Fe was associated with short-timescale (2–4 years) increases in precipitation, leading to a temporal decoupling between Fe and TOC. Our findings suggest that Fe concentrations do not increase uniformly with rising temperatures and increased precipitation, especially in regions where sulfur deposition has declined due to atmospheric recovery policies.

Zooplankton community composition of northern lakes is changing due to the interactive effects of climate change and recovery from acidification, yet limited data are available to assess these changes combined. Here, we built a database using archives of temperature, water chemistry and zooplankton data from 60 Scandinavian lakes that represent broad spatial and temporal gradients in key parameters: temperature, calcium (Ca), total phosphorus (TP), total organic carbon (TOC), and pH. Using machine learning techniques, we found that Ca was the most important determinant of the relative abundance of all zooplankton groups studied, while pH was second, and TOC third in importance. Further, we found that Ca is declining in almost all lakes, and we detected a critical Ca threshold in lake water of 1.3 mg L−1, below which the relative abundance of zooplankton shifts toward dominance of Holopedium gibberum and small cladocerans at the expense of Daphnia and copepods. Our findings suggest that low Ca concentrations may shape zooplankton communities, and that current trajectories of Ca decline could promote widespread changes in pelagic food webs as zooplankton are important trophic links from phytoplankton to fish and different zooplankton species play different roles in this context.

Global change may introduce fundamental alterations in phytoplankton biomass and community structure that can alter the productivity of northern lakes. In this study, we utilized Swedish and Finnish monitoring data from lakes that are spatially (135 lakes) and temporally (1995-2019, 110 lakes) extensive to assess how phytoplankton biomass (PB) of dominant phytoplankton groups related to changes in water temperature, pH and key nutrients [total phosphorus (TP), total nitrogen (TN), total organic carbon (TOC), iron (Fe)] along spatial (Fennoscandia) and temporal (25 years) gradients. Using a machine learning approach, we found that TP was the most important determinant of total PB and biomass of a specific species of Raphidophyceae - Gonyostomum semen - and Cyanobacteria (both typically with adverse impacts on food-webs and water quality) in spatial analyses, while Fe and pH were second in importance for G. semen and TN and pH were second and third in importance for Cyanobacteria. However, in temporal analyses, decreasing Fe and increasing pH and TOC were associated with a decrease in G. semen and an increase in Cyanobacteria. In addition, in many lakes increasing TOC seemed to have generated browning to an extent that significantly reduced PB. The identified discrepancy between the spatial and temporal results suggests that substitutions of data for space-for-time may not be adequate to characterize long-term effects of global change on phytoplankton. Further, we found that total PB exhibited contrasting temporal trends (increasing in northern- and decreasing in southern Fennoscandia), with the decline in total PB being more pronounced than the increase. Among phytoplankton, G. semen biomass showed the strongest decline, while cyanobacterial biomass showed the strongest increase over 25 years. Our findings suggest that progressing browning and changes in Fe and pH promote significant temporal changes in PB and shifts in phytoplankton community structures in northern lakes.

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.

Accounting for temporal changes in carbon dioxide (CO₂) effluxes from freshwaters remains a challenge for global and regional carbon budgets. Here, we synthesize 171 site-months of flux measurements of CO₂ based on the eddy covariance method from 13 lakes and reservoirs in the Northern Hemisphere, and quantify dynamics at multiple temporal scales. We found pronounced sub-annual variability in CO₂ flux at all sites. By accounting for diel variation, only 11% of site-months were net daily sinks of CO₂. Annual CO₂ emissions had an average of 25% (range 3%-58%) interannual variation. Similar to studies on streams, nighttime emissions regularly exceeded daytime emissions. Biophysical regulations of CO₂ flux variability were delineated through mutual information analysis. Sample analysis of CO₂ fluxes indicate the importance of continuous measurements. Better characterization of short- and long-term variability is necessary to understand and improve detection of temporal changes of CO₂ fluxes in response to natural and anthropogenic drivers. Our results indicate that existing global lake carbon budgets relying primarily on daytime measurements yield underestimates of net emissions.

1. Global changes are causing decreases in inorganic nitrogen (N) concentrations, increases in coloured dissolved organic carbon (DOC) concentrations, and decreases in dissolved inorganic N to total phosphorus ratios (DIN:TP) in northern lakes. The effects of these changes on phytoplankton and zooplankton biomass and the N:P recycling ratio of zooplankton remain unresolved.
2. In 33 Swedish headwater lakes across subarctic-to-boreal gradients with different levels of N deposition (low N in the north [Västerbotten, boreal; Abisko, subarctic] vs. high N in the south [Värmland, boreal; Jämtland, subarctic]), we measured water chemistry, phytoplankton biomass (chlorophyll-a [Chl-a], Chl-a:TP), seston mineral quality (C:P, N:P), as well as zooplankton biomass, community composition, and C:N:P stoichiometry. We estimated nutrient imbalances and the N:P recycling ratios of zooplankton using ecological stoichiometry models.
3. There was a large-scale gradient from low lake DIN and DIN:TP in the north to high DIN and DIN:TP in the south, with lower DIN:TP in lakes coinciding with higher DOC within each region. Lower lake DIN was associated with lower phytoplankton biomass (lower Chl-a:TP). Lower lake DIN:TP was associated with richer seston mineral quality (lower seston C:P and N:P) and higher zooplankton biomass.
4. Zooplankton community composition differed in the north vs. south, with a dominance of N-requiring calanoid copepods with high N:P in the north and P-requiring cladocerans with low N:P in the south. Also, greater differences in zooplankton community composition were found between subarctic regions (with lower DOC) than between boreal regions (with higher DOC), suggesting that increases in lake DOC and associated declines in lake DIN:TP reduce differences in zooplankton community composition.
5. The combination of lower lake DIN, higher lake DOC, and lower lake DIN:TP led to reduced zooplankton N:P recycling ratios, possibly by reducing seston N:P and/or by enhancing calanoid copepod dominance in the zooplankton community.
6. Our findings suggest that the combination of declining N deposition and increasing lake browning in northern high-latitude lakes will reduce phytoplankton biomass, but will concurrently enhance seston mineral quality and probably also zooplankton biomass and their recycling efficiency of P relative to N.

We used data from whole-lake studies to assess how changes in food quantity (phytoplankton biomass) and quality (phytoplankton community composition, seston C:P and N:P) with N fertilization affect zooplankton biomass, community composition and C:N:P stoichiometry, and their N:P recycling ratio along a gradient in lake DOC concentrations. We found that despite major differences in phytoplankton biomass with DOC (unimodal distributions, especially with N fertilization), no major differences in zooplankton biomass were detectable. Instead, phytoplankton to zooplankton biomass ratios were high, especially at intermediate DOC and after N fertilization, implying low trophic transfer efficiencies. An explanation for the observed low phytoplankton resource use, and biomass responses in zooplankton, was dominance of colony forming chlorophytes of reduced edibility at intermediate lake DOC, combined with reduced phytoplankton mineral quality (enhanced seston N:P) with N fertilization. N fertilization, however, increased zooplankton N:P recycling ratios, with largest impact at low DOC where phytoplankton benefitted from light sufficiently to cause enhanced seston N:P. Our results suggest that although N enrichment and increased phytoplankton biomass do not necessarily increase zooplankton biomass, bottom-up effects may still impact zooplankton and their N:P recycling ratio through promotion of phytoplankton species of low edibility and altered mineral quality.