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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.

Dietary uptake is key for transferring potentially toxic contaminants, such as mercury (Hg) and essential dietary nutrients, such as polyunsaturated fatty acids (PUFA), to consumers at higher trophic levels of aquatic food webs. We evaluated the role of diet sources for Hg bioaccumulation and PUFA retention in fish across lake food webs in seven Swedish lakes and two Chinese reservoirs. Fish total Hg (THg) and methyl-Hg (MeHg) differed greatly between the two countries: the Chinese fish contained less than 300 ng g⁻¹ dry weight (d.w.) THg with less than 50% as MeHg, versus the Swedish fishes which contained approximately 2000 ng g⁻¹ d.w. THg and nearly 100% as MeHg. Fatty acids enrichment of linoleic acids (LIN) were more prevalent in the Chinese fishes regardless of size (p < 0.05). Here we examined food web length, fish growth rates, and fatty acids patterns in relation to the quality of fish as a food source for both Hg and FA. Contrary to the expectation that biodilution of Hg throughout the food chain would explain these differences, a more complex picture emerged with high levels of Hg at the base of the food web in the Chinese reservoirs, a decoupling of fatty acid and Hg bioaccumulation, and a major role for both fish stocking and fish feed. It is hoped that this work will provide a nuanced picture of fish quality as a food source in different ecosystems.

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.

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.

1. Globally, lakes are warming and browning with ongoing climate change. These changes significantly impact a lake's biogeochemical properties and all organisms, including invertebrate consumers. The effects of these changes are essential to understand, especially during critical periods after and before the growing season, that is, autumn and spring, which can determine the composition of the invertebrate consumer community.
2. In this study, we used a large-scale experimental pond system to test the combined effect of warming (+3°C) and increased input of terrestrial and coloured dissolved organic carbon (gradient of 1.6–8.8 mg/L in the ambient and 1.6–9.3 mg/L in the warm)—which causes browning—on zooplankton and benthic macroinvertebrate biomass and composition during the autumn and the following spring.
3. Total zooplankton biomass decreased with warming and increased with browning, while total zoobenthos did not respond to either treatment. Warming and browning throughout the autumn had no overall interactive effects on zooplankton or zoobenthos. Autumnal warming decreased total pelagic consumer biomass, caused by a decrease in both Rotifera and Copepoda. In contrast, there was no effect on overall benthic consumer biomass, with only Asellus sp. biomass showing a negative response to warming. An autumnal increase in dissolved organic carbon led to increased total pelagic consumer biomass, which was related to increases in Daphnia sp. biomass but did not affect zoobenthos biomass. While we expected zooplankton and zoobenthos biomass to follow responses in primary and bacterial production to treatments, we did not find any relationship between consumer groups and these estimates of resource production.
4. Our results suggest that consumer responses to warming and browning during autumn may lead to less overarching general changes in consumer biomass, and responses are mostly taxon-specific.
5. This study gives novel insights into the effects of warming and browning on consumer biomass during autumn and spring and increases the understanding of the effects of climate change on invertebrate community biomass in the different habitats.

Aquatic microbial responses to changes in the amount and composition of dissolved organic carbon (DOC) are of fundamental ecological and biogeochemical importance. Parallel factor (PARAFAC) analysis of excitation–emission fluorescence spectra is a common tool to characterize DOC, yet its ability to predict bacterial production (BP), bacterial respiration (BR), and bacterial growth efficiency (BGE) vary widely, potentially because inorganic nutrient limitation decouples microbial processes from their dependence on DOC composition. We used 28-d bioassays with water from 19 lakes, streams, and rivers in northern Sweden to test how much the links between bacterial metabolism and fluorescence PARAFAC components depend on experimental additions of inorganic nutrients. We found a significant interaction effect between nutrient addition and fluorescence on carbon-specific BP, and weak evidence for influence on BGE by the same interaction (p = 0.1), but no corresponding interaction effect on BR. A practical implication of this interaction was that fluorescence components could explain more than twice as much of the variability in carbon-specific BP (R² = 0.90) and BGE (R² = 0.70) after nitrogen and phosphorus addition, compared with control incubations. Our results suggest that an increased supply of labile DOC relative to ambient phosphorus and nitrogen induces gradually larger degrees of nutrient limitation of BP, which in turn decouple BP and BGE from fluorescence signals. Thus, while fluorescence does contain precise information about the degree to which DOC can support microbial processes, this information may be hidden in field studies due to nutrient limitation of bacterial metabolism.

In recent years, unexplained declines in lake total phosphorus (TP) concentrations have been observed at northern latitudes (> 42°N latitude) where most of the world's lakes are found. We compiled data from 389 lakes in Fennoscandia and eastern North America to investigate the effects of climate on lake TP concentrations. Synchrony in year-to-year variability is an indicator of climatic influences on lake TP, because other major influences on nutrients (e.g., land use change) are not likely to affect all lakes in the same year. We identified significant synchrony in lake TP both within and among different geographic regions. Using a bootstrapped random forest analysis, we identified winter temperature as the most important factor controlling annual TP, followed by summer precipitation. In Fennoscandia, TP was negatively correlated with the winter East Atlantic Pattern, which is associated with regionally warmer winters. Our results suggest that, in the absence of other overriding factors, lake TP and productivity may decline with continued winter warming 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.

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.