A key issue to understanding the transformations of terrestrial organic carbon in the ocean is to disentangle the latter from marine-produced organic matter. We applied a multiple stable isotope approach using delta(34)S and delta(13)C isotope signatures from estuarine dissolved organic matter (DOM), enabling us to constrain the contribution of terrestrial-derived DOM in an estuarine gradient of the northern Baltic Sea. The stable isotope signatures for dissolved organic sulfur (delta(34)S(DOS)) have twice the range between terrestrial and marine end members compared to the stable isotope signatures for dissolved organic carbon (delta(13)C(DOC)); hence, the share of terrestrial DOM in the total estuarine DOM can be calculated more precisely. DOM samples from the water column were collected using ultrafiltration on board the German RV Maria S Merian during a winter cruise, in the Bothnian Bay, Bothnian Sea, and Baltic proper. We calculated the terrestrial fraction of the estuarine DOC (DOC(ter)) from both delta(13)C(DOC) and delta(34)S(DOS) signatures and applying fixed C: S ratios for riverine and marine end members to convert S isotope signatures into DOC concentrations. The delta(34)S(DOS) signature of the riverine end member was +7.02 parts per thousand, and the mean signatures from Bothnian Bay, Bothnian Sea, and Baltic proper were +10.27, +12.51, and +13.67 parts per thousand, respectively, showing an increasing marine signal southwards (d34SDOS marine end member = 18.1 parts per thousand). These signatures indicate that 87%, 75%, and 67%, respectively, of the water column DOC is of terrestrial origin (DOC(ter)) in these basins. Comparing the fractions of DOC(ter) in each basin-that are still based on few winter values only-with the annual river input of DOC, it appears that the turnover time for DOC(ter) in the Gulf of Bothnia is much shorter than the hydraulic turnover time, suggesting that high-latitude estuaries might be efficient sinks for DOC(ter).

Abstract The contribution of terrestrially derived carbon to micro-crustacean zooplankton biomass (i.e., allochthony) has been previously studied in lakes, reservoirs, and estuaries, but little is known about zooplankton allochthony in rivers. In lacustrine environments, allochthony is regulated by distinct selective feeding behavior of different taxa. However, we hypothesized that restricted possibility for selective grazing in turbulent environments such as rivers would decouple zooplankton from specific microbial and algal food resources, such that their allochthony would mirror the terrestrial contribution to the surrounding bulk particle pool. We tested this idea by analyzing allochthony in 13 widely distributed Swedish rivers, using a dual-isotope mixing model. Zooplankton biomasses were generally low, and allochthony in different micro-crustacean groups (Cladocera, Cyclopoida, Calanoida) varied from 2% to 77%. As predicted, there were no correlations between allochthony and variables indicating the supply of algal and microbial food resources, such as chlorophyll a and bacterial production. Instead, the allochthony was generally similar to the share allochthonous contribution in bulk particulate organic matter, with relationships close to the 1 : 1 line. The zooplankton community allochthony was strongly regulated by the ecosystem metabolic balance between production and respiration, which in turn was dependent upon the ratio between total autochthonous organic carbon concentrations and water color. Our study for the first time shows that micro-crustacean allochthony is regulated differently in rivers compared to in lacustrine systems, and points to inefficient support of zooplankton biomass by algal resources in turbulent waters.

The ecology and biogeochemistry of lakes in the subarctic region are particularly sensitive to changes in the abundance and optical properties of dissolved organic matter (DOM). External input of colored DOM to these lakes is an extensively researched topic, but little is known about potential reciprocal feedbacks between the optical properties of DOM and internal microbial processes in the water. We performed 28-day dark laboratory incubation trials on water from 101 subarctic tundra lakes in northern Sweden, measuring the microbial decay of DOM and the resulting dynamics in colored (CDOM) and fluorescent (FDOM) DOM components. While losses in dissolved oxygen during the incubations corresponded to a 20% decrease in mean DOM, conversely the mean CDOM and total FDOM increased by 22% and 30%, respectively. However, the patterns in microbial transformation of the DOM were not the same in all lakes. Notably, along the gradient of increasing ambient CDOM (water brownness), the lakes showed decreased microbial production of protein-like fluorescence, lowered DOM turnover rates and decreasing bacterial growth per unit of DOM. These trends indicate that browning of subarctic lakes systematically change the way that bacteria interact with the ambient DOM pool. Our study underscores that there is no unidirectional causal link between microbial processes and DOM optical properties, but rather reciprocal dependence between the two.

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.

Over the past decade, it has been Shown that unproductive lakes worldwide are net heterotrophic because bacterial respiration of allochthonous. organic carbon (AOC) makes community respiration exceed primary production. Net heterotrophy means that aquatic systems are net sources of CO2 to the atmosphere but also that bacterial utilization of AOC increases bacterioplankton production (BP) and bacterial uptake of limiting inorganic nutrients at the expense of phytoplankton production (PP). We studied 15 unproductive lakes in northern Sweden with dissolved organic carbon concentrations between 3 and 22 mg L-1. We found a highly significant negative relationship between the degree of heterotrophy and total pelagic energy mobilization (PP + BP based on AOC) per unit of limiting nutrient. We suggest that this is because the high cell phosphorous (P) requirement of bacteria makes energy mobilization per P unit considerably lower in bacterioplankton than in phytoplankton. We also suggest that the productivity of the entire pelagic ecosystem is determined by the availability of inorganic nutrients and AOC and by whether nutrients are allocated to BP or PP.

We compared the stable carbon isotopic composition (delta(13)C) of crustacean zooplankton with that of potential carbon sources in 15 lakes in northern Sweden with different dissolved organic carbon (DOC) concentrations (2-9 mg L-1) to test the hypothesis that zooplankton depended more on allochthonous carbon in humic lakes than in clear-water lakes. Based on delta(13)C signature, we found that the pool of organic matter in the lakes was dominated by carbon of allochthonous origin over the whole DOC gradient. Zooplankton were generally depleted in C-13 compared to organic matter in the catchment, particulate organic matter in the lake water, and shallow surface sediment. However, the isotopic composition of zooplankton could not be explained without a significant contribution from both allochthonous and autochthonous carbon sources in all lakes. The relative importance of these two carbon sources did not relate to the concentration of, or proportion between, allochthonous and autochthonous organic carbon in the water. Instead, the proportion between allochthonous and autochthonous carbon in the crustacean zooplankton was consistent with a rather conservative use of the energy mobilized by bacterioplankton and phytoplankton in the lakes.

Metabolism is one of the most fundamental ecosystem processes, but the drivers of variation in metabolic rates among lakes dominated by benthic primary producers remain poorly constrained. Here, we report the magnitudes and potential drivers of whole-lake metabolism across 43 Swedish arctic–alpine lakes, based on the free-water diel oxygen technique with sondes deployed during the open-water season near the surface and bottom of the lakes. Gross primary production (GPP) and ecosystem respiration (R) were strongly coupled and ranged from 0.06 to 0.45 mg and 0.05 to 0.43 mg L⁻¹ d⁻¹ among lakes. On average, GPP and R decreased eightfold from relatively shallow to deep lakes (mean depth 0.5–10.9 m) and twofold from concave to convex lakes (mean depth: maximum depth 0.2–0.5). We attribute this to light limitation and shape-specific sensitivity of benthic GPP to disturbance by lake ice. Net ecosystem production (GPP-R) ranged from −0.09 to 0.14 mg L⁻¹ d⁻¹ and switched, on average, from positive to negative towards deeper lakes and lakes richer in dissolved organic carbon (DOC; 0.5–7.4 mg DOC L⁻¹). Uncertainties in metabolism estimates were high (around one and three times mean R and GPP), especially in deep lakes with low insulation and diurnally variable wind speed. Our results confirm the role of DOC in stimulating net heterotrophy and highlight novel effects of lake shape on productivity in benthic-dominated lake ecosystems and its response to changes in lake ice cover.

The seasonality of gross primary production (GPP) in streams is driven by multiple physical and chemical factors, yet incident light is often thought to be most important. In Arctic tundra streams, however, light is available in saturating amounts throughout the summer, but sharp declines in nutrient supply during the terrestrial growing season may constrain aquatic productivity. Given the opposing seasonality of these drivers, we hypothesized that "shoulder seasons"-spring and autumn-represent critical time windows when light and nutrients align to optimize rates of stream productivity in the Arctic. To test this, we measured annual patterns of GPP and biofilm accumulation in eight streams in Arctic Sweden. We found that the aquatic growing season length differed by 4 months across streams and was determined largely by the timing of ice-off in spring. During the growing season, temporal variability in GPP for nitrogen (N) poor streams was correlated with inorganic N concentration, while in more N-rich streams GPP was instead linked to changes in phosphorus and light. Annual GPP varied ninefold among streams and was enhanced by N availability, the length of ice-free period, and low flood frequency. Finally, network scale estimates of GPP highlight the overall significance of the shoulder seasons, which accounted for 48% of annual productivity. We suggest that the timing of ice off and nutrient supply from land interact to regulate the annual metabolic regimes of nutrient poor, Arctic streams, leading to unexpected peaks in productivity that are offset from the terrestrial growing season.

Global warming is enhancing the mobilization of organic carbon (C) from Arctic soils into streams, where it can be mineralized to CO₂ and released to the atmosphere. Abiotic photo‐oxidation might drive C mineralization, but this process has not been quantitatively integrated with biological processes that also influence CO₂ dynamics in aquatic ecosystems. We measured CO₂ concentrations and the isotopic composition of dissolved inorganic C (δ¹³C_DIC) at diel resolution in two Arctic streams, and coupled this with whole‐system metabolism estimates to assess the effect of biotic and abiotic processes on stream C dynamics. CO₂ concentrations consistently decreased from night to day, a pattern counter to the hypothesis that photo‐oxidation is the dominant source of CO₂. Instead, the observed decrease in CO₂ during daytime was explained by photosynthetic rates, which were strongly correlated with diurnal changes in δ¹³C_DIC values. However, on days when modeled photosynthetic rates were near zero, there was still a significant diel change in δ¹³C_DIC values, suggesting that metabolic estimates are partly masked by O₂ consumption from photo‐oxidation. Our results suggest that 6–12 mmol CO₂‐C m⁻² d⁻¹ may be generated from photo‐oxidation, a range that corresponds well to previous laboratory measurements. Moreover, ecosystem respiration rates were 10 times greater than published photo‐oxidation rates for these Arctic streams, and accounted for 33–80% of total CO₂ evasion. Our results suggest that metabolic activity is the dominant process for CO₂ production in Arctic streams. Thus, future aquatic CO₂ emissions may depend on how biotic processes respond to the ongoing environmental change.

The effects of climatic variables on lake‐water total organic carbon (TOC) concentrations and benthic and pelagic primary producers during the past 45 yr were assessed using the sediment records of two subarctic lakes, one with mires and one without mires connected to the lake. The lake with a mire showed large and synchronous changes in the planktonic to benthic (P : B) ratio of diatoms and concentrations of TOC inferred from nearinfrared spectroscopy. During periods of warm temperatures, high precipitation, and long ice‐free conditions, we inferred high TOC in the lake, and the diatom community was dominated by planktonic species. The stable carbon isotopic (Δ¹³C) values of sediment organic matter were negatively correlated with inferred TOC concentration and P :B ratio. We suggest that the changes in TOC and P : B ratio were a result of changing climate, permafrost degradation, and related changes in the catchment. Terrestrial organic matter, by its strong effect on the penetration of light through the lake water, possibly affected the habitats available for benthic photosynthesis and thus the Δ13C of the sediment organic matter. The large changes in recent times may also be because of unusually long ice‐free periods, warmer temperatures, and other associated limnological changes. The lake with no mire next to the lake showed only minor changes in lake‐water TOC during the same period and P :B ratio remained almost constant until the past 5 yr, when the P :B ratio increased rapidly. The observed changes in P :B ratio within this lake may be because of complex interactions of several climate‐related variables.

The Arctic Ocean is more susceptible to ocean acidification than other marine environments due to its weaker buffering capacity, while its cold surface water with relatively low salinity promotes atmospheric CO2 uptake. We studied how sea-ice microbial communities in the central Arctic Ocean may be affected by changes in the carbonate system expected as a consequence of ocean acidification. In a series of four experiments during late summer 2018 aboard the icebreaker Oden, we addressed microbial growth, production of dissolved organic carbon (DOC) and extracellular polymeric substances (EPS), photosynthetic activity, and bacterial assemblage structure as sea-ice microbial communities were exposed to elevated partial pressures of CO2 (pCO2). We incubated intact, bottom ice-core sections and dislodged, under-ice algal aggregates (dominated by Melosira arctica) in separate experiments under approximately 400, 650, 1000, and 2000μatm pCO2 for 10 d under different nutrient regimes. The results indicate that the growth of sea-ice algae and bacteria was unaffected by these higher pCO2 levels, and concentrations of DOC and EPS were unaffected by a shifted inorganic C/N balance, resulting from the CO2 enrichment. These central Arctic sea-ice microbial communities thus appear to be largely insensitive to short-term pCO2 perturbations. Given the natural, seasonally driven fluctuations in the carbonate system of sea ice, its resident microorganisms may be sufficiently tolerant of large variations in pCO2 and thus less vulnerable than pelagic communities to the impacts of ocean acidification, increasing the ecological importance of sea-ice microorganisms even as the loss of Arctic sea ice continues.