We measured sediment production of carbon dioxide (CO(2)) and methane (CH(4)) and the net flux of CO(2) across the surfaces of 15 boreal and subarctic lakes of different humic contents. Sediment respiration measurements were made in situ under ambient light conditions. The flux of CO(2) between sediment and water varied between an uptake of 53 and an efflux of 182 mg C m(-2) day(-1) from the sediments. The mean respiration rate for sediments in contact with the upper mixed layer (SedR) was positively correlated to dissolved organic carbon (DOC) concentration in the water (r(2) = 0.61). The net flux of CO(2) across the lake surface [net ecosystem exchange (NEE)] was also closely correlated to DOC concentration in the upper mixed layer (r(2) = 0.73). The respiration in the water column was generally 10-fold higher per unit lake area compared to sediment respiration. Lakes with DOC concentrations <5.6 mg L(-1) had net consumption of CO(2) in the sediments, which we ascribe to benthic primary production. Only lakes with very low DOC concentrations were net autotrophic (<2.6 mg L(-1)) due to the dominance of dissolved allochthonous organic carbon in the water as an energy source for aquatic organisms. In addition to previous findings of allochthonous organic matter as an important driver of heterotrophic metabolism in the water column of lakes, this study suggests that sediment metabolism is also highly dependent on allochthonous carbon sources.
Inorganic nutrient availability and temperature are recognized as major regulators of organic carbon processing by aquatic bacteria, but little is known about how these two factors interact to control bacterial metabolic processes. We manipulated the temperature of boreal humic stream water samples within 0–25°C and measured bacterial production (BP) and respiration (BR) with and without inorganic nitrogen + phosphorus addition. Both BP and BR increased exponentially with temperature in all experiments, with Q 10 values varying between 1.2 and 2.4. The bacterial growth efficiency (BGE) showed strong negative relationships with temperature in nutrient-enriched samples and in natural stream water where community-level BP and BR were not limited by nutrients. However, there were no relationships between BGE and temperature in samples where BP and BR were significantly constrained by the inorganic nutrient availability. The results suggest that metabolic responses of aquatic bacterial communities to temperature variations can be strongly dependent on whether the bacterial metabolism is limited by inorganic nutrients or not. Such responses can have consequences for both the carbon flux through aquatic food webs and for the flux of CO2 from aquatic systems to the atmosphere.
Ecology Letters (2010) Abstract Carbon of terrestrial origin often makes up a significant share of consumer biomass in unproductive lake ecosystems. However, the mechanisms for terrestrial support of lake secondary production are largely unclear. By using a modelling approach, we show that terrestrial export of dissolved labile low molecular weight carbon (LMWC) compounds supported 80% (34-95%), 54% (19-90%) and 23% (7-45%) of the secondary production by bacteria, protozoa and metazoa, respectively, in a 7-km(2) boreal lake (conservative to liberal estimates in brackets). Bacterial growth on LMWC was of similar magnitude as that of primary production (PP), and grazing on bacteria effectively channelled the LMWC carbon to higher trophic levels. We suggest that rapid turnover of forest LMWC pools enables continuous export of fresh photosynthates and other labile metabolites to aquatic systems, and that substantial transfer of LMWC from terrestrial sources to lake consumers can occur within a few days. Sequestration of LMWC of terrestrial origin, thus, helps explain high shares of terrestrial carbon in lake organisms and implies that lake food webs can be closely dependent on recent terrestrial PP.
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.
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.
Phytoplankton play a key role in supporting aquatic food webs. However, the effects of ongoing large-scale changes in the concentrations and stoichiometry of important biological compounds [dissolved inorganic N (DIN), total phosphorus (TP), dissolved organic carbon (DOC) and DIN:TP] on the development and nutritional quality of phytoplankton for higher trophic levels are unclear. We conducted lake studies and in situ bioassay experiments in two Swedish mountain regions [Abisko (north) and Jamtland (south)] with different N deposition and where lakes in each region were distributed along a similar gradient in lake DOC (2-7 mg L-1) to assess whether differences in nutrients, DOC and DIN:TP induced differences in phytoplankton quantity [chlorophyll a (Chl-a) and seston carbon (C)] and quality [seston C:N:P stoichiometry and fatty acid (FA) composition]. Using long-term monitoring data from lakes in these two mountain regions, we found declining long-term trends in N deposition and lake DIN and total TP concentrations, but not in lake DIN:TP. Lakes in Abisko received lower N deposition and had lower DIN:TP than those in Jamtland. Phytoplankton was N- to NP-limited in Abisko lakes but NP dual-limited in Jamtland lakes. The N fertilization effects induced by higher DIN:TP were weak on phytoplankton quantity but strong on phytoplankton quality. The phytoplankton had lower eicosapentaenoic acid (EPA) content and higher P content (lower seston C:P) in Abisko compared to in Jamtland. In addition, the quality of the DOC (as indicated by its aromaticity and SUVA) influenced not only the light conditions and the seston C:P ratios, but also the FA composition. We found higher bacteria FA concentrations in seston in Abisko than in Jamtland, despite lower amounts of FA of terrestrial origin in Abisko. Our findings suggest that declining N deposition and enhanced colored terrestrial C loadings leads to lower nutritional quality of basal resources for higher consumers in mountain lakes.
Lake sampling and in situ nutrient enrichment enclosure experiments with nitrogen (N) and phosphorus (P) were conducted in unproductive Swedish lakes along a gradient of increasing atmospheric N-deposition. The regional and seasonal patterns of nutrient limitation of phytoplankton were clearly related to the amounts of N-deposition and N-inputs the lakes received. In areas of low N-deposition in northern Sweden, N-limitation of phytoplankton was evident throughout the summer season due to high catchment N-retention and very low dissolved inorganic N (DIN) inputs during the early summer. High N-deposition in the south was accompanied by high lake DIN-concentrations during the early summer and subsequent P-limitation of phytoplankton. However, P-limitation did not persist over the summer and, as a consequence of a declining DIN-pool, the lakes switched to dual- and co-limitation by N and P, and then to N-limitation. Generally, the lakes were N-limited rather than P-limited during the summer. We conclude that N-limitation is probably a natural state of the unproductive lakes studied, but P-limitation of variable intensity and duration has been induced by elevated atmospheric N-deposition.
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.
Northern lakes are experiencing widespread increases in dissolved organic carbon (DOC) that are likely to lead to changes in pelagic phytoplankton biomass. Pelagic phytoplankton biomass responds to trade-offs between light and nutrient availability. However, the influence of DOC light absorbing properties and carbon–nutrient stoichiometry on phytoplankton biomass across seasonal or spatial gradients has not been assessed. Here, we analyzed data from almost 5000 lakes to examine how the carbon–phytoplankton biomass relationship is influenced by seasonal changes in light availability, DOC light absorbing properties (carbon-specific visual absorbance, SVA420), and DOC–nutrient [total nitrogen (TN) and total phosphorus (TP)] stoichiometry, using TOC as a proxy for DOC. We found evidence for trade-offs between light and nutrient availability in the relationship between DOC and phytoplankton biomass [chlorophyll (chl)-a], with the shape of the relationship varying with season. A clear unimodal relationship was found only in the fall, particularly in the subsets of lakes with the highest TOC:TP. Observed trends of increasing TOC:TP and decreasing TOC:TN suggest that the effects of future browning will be contingent on future changes in carbon–nutrient stoichiometry. If browning continues, phytoplankton biomass will likely increase in most northern lakes, with increases of up to 76% for a 1.7 mg L−1 increase in DOC expected in subarctic regions, where DOC, SVA420, DOC:TN, and DOC:TP are all low. In boreal regions with higher DOC and higher SVA420, and thus lower light availability, lakes may experience only moderate increases or even decreases in phytoplankton biomass with future browning.
Concentrations of dissolved organic carbon (DOC) have increased recently in many lakes at high latitudes in North America and Europe, but it is unclear what effect this will have on the identity of the limiting nutrient for phytoplankton [nitrogen (N) vs. phosphorus (P)]. Identifying the effect of changing DOC on phytoplankton nutrient limitation is complicated by spatial covariation between atmospheric N deposition and increasing DOC in areas where lake browning occurs. We conducted nutrient-limitation assays in 27 lakes from three sites along gradients of climate and atmospheric N deposition in Sweden. Within each site, lakes were selected to represent the range of DOC concentrations. We also conducted statistical analyses of large-scale lake survey data (n = 4768 lakes divided into 47 regions) to investigate relationships between DOC and nutrient stoichiometry while controlling for differences in N deposition. Our findings confirmed that most lakes were dual-limited by both N and P in the south, whereas northern lakes were primarily N-limited. Throughout Sweden the ratio of dissolved inorganic nitrogen (DIN) to total phosphorus (TP) declined with increasing DOC in most regions, suggesting that browner lakes are more likely to be N limited. These results were not supported by our nutrient limitation assays, which identified no relationship between DOC and relative strength of limitation by N or P. Increased DOC also resulted in significant increases in both total and inorganic N and P fractions, suggesting that other factors such as light limitation or increased top-down control become more important as DOC increases.
Bioavailable phosphorus (BAP) concentrations were determined nine times between April and October 2010 in two humic boreal headwater streams draining forest-and mire-dominated catchments. BAP was analyzed in a bioassay in which natural P-limited bacterioplankton grew with natural P as the sole P source. In both streams, approximately 90% of the BAP occurred as dissolved species (passing a 0.2-mu m filter), consisting partly of low-molecular-weight forms (passing a filter with nominal cutoff at 1 kDa) and partly of high-molecular-weight forms (passing a 0.2-mu m filter but not a 1-kDa filter). Concentrations of total dissolved BAP varied between 1 mu g L-1 and 14 mu g L-1, with the highest values in the middle of the summer. Compared to the forest stream, BAP concentrations were generally higher in the mire stream, where it occasionally amounted to nearly 50% of total P. Molybdate reactive phosphorus overestimated BAP considerably. Most of the BAP was in forms other than free orthophosphate. Temporal BAP variations showed no relationships with dissolved organic carbon (C) or iron but were positively related to air temperature and negatively related to the absorbance ratio (a254 : a365) of organic compounds in the water, indicating connections between terrestrial export of BAP and temperature-dependent terrestrial C metabolism. Concentrations of BAP can relieve stream bacteria from P limitation, and a significant share of BAP exported to streams can reach and be used in downstream lakes.
Bacterial utilization of allochthonous organic carbon and the subsequent fate of the bacterial production in the pelagic food web were estimated in Lake Örträsket, a large humic lake in northern Sweden. Bacterial production relied mainly on allochthonous DOC and exceeded primary production in the epilimnion. Bacterial productivity was clearly stimulated during high flow episodes, increasing the input of bacteria degradable organic material to the lake. Bacterioplankton were exploited mainly by mixotrophic flagellates which probably used bacteria as a source of carbon, phosphorus and nitrogen. The possibly extremely low availability of inorganic P during periods with high bacterial production may have allowed the mixotrophs to outcompete obligate autotrophs and help them become dominant phytoplankters during large parts of the summer. The results from Lake Örträsket indicated that the total production depended on bacterial energy mobilization from allochthonous organic carbon compounds and that heterotrophic mobilized energy was linked via mixotrophs to higher levels in the food chain.
Enrichment experiments with P and N were conducted in humic Lake Örträsket in northern Sweden. The composition of the microplankton community showed a dominance by bacterioplankton, followed by mixotrophic sind potentially mixotrophic phytoplankton, heterotrophic nanoflagellates, and autotrophic phytoplankton. Bacterioplankton was P limited for most of the ice-free period, and phytoplankton biomass and primary production mostly increased after enrichment with N, but not with P. The dominant group of phytoplankton, the mixotrophic flagellates, was stimulated by N bur not by P, while obligate autotrophic species were stimulated only by P+N. It is suggested that N limitation in mixotrophic species is induced by grazing of P-rich bacteria. The results suggest that primary productivity in humic lakes can be limited by N and indicate the importance of phagocytosis as a means of nutrition in phytoplankton. A link is suggested to exist in humic lakes whereby heterotrophic bacterioplankton, which use humic compounds as their principal energy source, can transfer energy and nutrients to potentially autotrophic organisms, with subsequent utilization by other components of the food web.
We compared terrestrial net primary production (NPP) and terrestrial export of dissolved organic carbon (DOC) with lake water heterotrophic bacterial activity in 12 headwater lake catchments along an altitude gradient in subarctic Sweden. Modelled NPP declined strongly with altitude and annual air temperature decreases along the altitude gradient (6ºC between the warmest and the coldest catchment). Estimated terrestrial DOC export to the lakes was closely correlated to NPP. Heterotrophic bacterial production (BP) and respiration (BR) were mainly based on terrestrial organic carbon and strongly correlated with the terrestrial DOC export. Excess respiration over PP of the pelagic system was similar to net emission of CO2 in the lakes. BR and CO2 emission made up considerably higher shares of the terrestrial DOC input in warm lakes than in cold lakes, implying that respiration and the degree of net heterotrophy in the lakes were dependant not only on terrestrial export of DOC, but also on characteristics in the lakes which changed along the gradient and affected the bacterial metabolization of allochthonous DOC. The study showed close links between terrestrial primary production, terrestrial DOC export and bacterial activity in lakes and how these relationships were dependant on air temperature. Increases in air temperature in high latitude unproductive systems might have considerable consequences for lake water productivity and release of CO2 to the atmosphere, which are ultimately determined by terrestrial primary production.
1. Air temperature will probably have pronounced effects on the composition of plankton communities in northern lake ecosystems, either via indirect effects on the export of essential elements from catchments or through direct effects of water temperature and the ice-free period on the behaviour of planktonic organisms.
2. We assessed the role of temperature by comparing planktonic communities in 15 lakes along a 6 °C air temperature gradient in subarctic Sweden.
3. We found that the biomass of phytoplankton, bacterioplankton and the total planktonic biomass were positively related to air temperature, probably as a result of climatic controls on the export of nitrogen from the catchment (which affects phytoplankton biomass) and dissolved organic carbon (affecting bacterioplankton biomass).
4. The structure of the zooplankton community, and top down effects on phytoplankton, were apparently not related to temperature but mainly to trophic interactions ultimately dependent on the presence of fish in the lakes.
5. Our results suggest that air temperature regimes and long-term warming can have strong effects on the planktonic biomass in high latitude lakes. Effects of temperature on the structure of the planktonic community might be less evident unless warming permits the invasion of fish into previous fishless lakes.
Ecology Letters (2012) Abstract A majority of the worlds lakes are supersaturated with respect to carbon dioxide (CO2). By experimental manipulation of the CO2 concentration in supersaturated boreal lakes, we demonstrate that phytoplankton primary production was up to 10 times higher in supersaturated lake water in comparison with water with CO2 at equilibrium concentrations and that CO2, together with nutrients, explained most of the variation in pelagic primary production and phytoplankton biomass over a wide variety of unproductive lakes. These results suggest that phytoplankton can be co-limited by CO2 and nutrients in unproductive lakes. As import of terrestrial organic carbon and its subsequent microbial mineralisation in lakes is a driving force of CO2-supersaturation our results suggest that lake productivity and carbon cycling may respond to variations in terrestrial organic carbon export, (e.g. caused by land use or climate change) in ways not described before.
The sedimentation of organic, amorphous oxides, crystalline oxides and crystalline silicate species of Fe and Al was investigated in humic Lake Ortrasket. The covariation with C, N and P sedimentation also was studied. The results showed a strong temporal variation in the occurrence of different Fe species. During high discharge, such as in spring, Fe mostly occurred as crystalline Fe-silicate species, while the amorphous form dominated at other times of the year. Al generally sedimented as crystalline Al-silicate species (95%). Amorphous Fe species showed a strong relation with organic C and N, suggesting an association between organic material and amorphous Fe oxides. P was not as strongly related to amorphous Fe, probably as a result of apatite or clay bound-P species sedimentation.
In this paper, we assess the extent to which the export of terrestrially fixed carbon to aquatic systems and the aquatic metabolism of this carbon affect the overall accumulation of organic carbon in a boreal catchment. We estimated the contribution of stocks and processes in aquatic environments to the carbon balance of a boreal catchment in northern Sweden. We used published data concerning the net ecosystem exchange (NEE) of CO2 in terrestrial environments, and calculations of loss of terrestrial carbon to surface water and the turnover of terrestrial carbon in aquatic systems. The NEE of terrestrial environments was estimated to be 139 g C/m2 of catchment area per year. The export of terrestrially fixed carbon to aquatic systems was 8.6 g C/m2/yr, resulting in a net accumulation of organic carbon in terrestrial systems of 131 g C/m2/yr. Almost 45% of the terrestrial export was mineralized in streams and lakes and evaded as CO2, while most of the remaining (approximately 55%) terrestrial export was transported to the sea as organic carbon or as dissolved inorganic carbon emanating from soil respiration. The sedimentation of organic carbon and input of organic carbon via aquatic primary production were insignificant when compared to the mineralization and river transport of terrestrial organic carbon. Aquatic fluxes were small compared to the terrestrial NEE, which we consider to be largely a consequence of the studied catchment being subject to intensive forestry resulting in a large annual accumulation of carbon in growing tree biomass.
Lake Örträsket is a large (7.3 km2) and deep (64 m) humic (DOC 10 mg-L-1) lake in northern Sweden. Sediment trap data (gross sedimentation) and sediment core data (net sedimentation) has been used to calculate sedimentation and mineralisation of carbon, nitrogen, phosphorus, iron, aluminium and total particulate matter (TPM). Yearly gross sedimentation of nitrogen and organic carbon was higher than yearly net sedimentation caused by an annual sediment mineralisation of 54 % and 32 % of gross sedimentation, respectively. Annual gross sedimentation of phosphorus, iron, aluminium and TPM equalled annual net sedimentation and no significant sediment mineralisation of these elements was calculated. The annual retention (input-output) of organic carbon and nitrogen approximately equalled net sedimentation. Sediment mineralisation of nitrogen resulted in an accumulation of nitrate in the water column of the lake during periods of lake stratification. Approximately 2/3 of the nitrogen mineralised in and lost from the sediment was transported out of the lake (as nitrate) via the outlet during turnover in spring and autumn. Remaining 1/3 was assumed to be lost through sediment denitrification, equalling a denitrification rate of about 1 mgN-m-2-y-1.
Partial pressure (pCO(2)) and flux to the atmosphere of carbon dioxide (CO2) were studied in northern alpine and forest lakes along a gradient of dissolved organic carbon (DOC) content (0.4-9.9 mg L-1). Sixteen lakes were each sampled three times over the course of the ice-free season, and an additional 35 lakes were sampled once at midsummer. pCO(2) data were acquired in the field by a headspace equilibration technique. Most lakes were supersaturated with CO2 along the entire DOC gradient, with relatively small seasonal differences. pCO(2) was positively correlated to DOC content, reflecting a close dependence between allochthonous DOC in-put and heterotrophic respiration in the lakes. Fluxes of CO2 to the atmosphere were estimated from the pCO(2) measurements. Benthic respiration was indicated to be important for CO2 emission in lakes with high DOC concentrations. In lakes with low DOC concentrations, pelagic mineralization alone was sufficient to account for a large part of the estimated fluxes.
Organic carbon mineralization was studied. in a large humic lake (Lake Örträsket) in northern Sweden during a well-defined summer stratification period following high water flow during snowmelt. Several independent methods including plankton counts, measurements of bacterioplankton and phytoplankton production, stable isotope monitoring, sediment trapping, and mass balance calculations were used. Total organic carbon mineralization showed a summer mean of 0.3 g C m(-2) d(-1) and was partitioned about equally between water and sediment. In the water column, organic matter was mineralized by bacteria (60%) and protozoan and metazoan zooplankton (30%), as well as by photooxidation (10%). Most of the mineralized organic carbon was of allochthonous origin. Primary production in the lake contributed at most 5% of the total organic carbon input and about 20% of the total organic carbon mineralization. Total carbon mineralization in. the epilimnion and metalimnion agreed well with an estimate of CO2 evasion from the stratified lake, while CO2 accumulation in the hypolimnion matched the O-2 consumption and resulted in a very negative delta C-13 of DIC before autumn overturn (-23 parts per thousand). Isotopic compositions of DIC and POC confirmed the dominant influence of terrestrial organic input on the cycling of both organic and inorganic carbon in the lake.
Low molecular weight organic carbon compounds are potentially important carbon and energy substrates to heterotrophic production in the aquatic environment. We studied the occurrence of dissolved free amino acids (AA), monosaccharides (CHO), and carboxylic acids (CA) in the subarctic Lake Diktar-Erik. The lake is unproductive with slightly humic water, and receives water via one major inlet stream draining a birch forest environment. The concentration of dissolved organic carbon (DOC) in the inlet stream was strongly correlated with the discharge. This relationship changed from season to season, indicating changes in the sources of the DOC entering the stream. AA and CHO each accounted for an average of less than 0.5% of the DOC. After high discharge events during the ice-free period, AA and CHO occurred in especially high concentrations. CA occurred in higher concentrations during the ice-free period, when it generally accounted for 20–30% of the DOC pool. The CA content relative to the total DOC pool was strongly inversely correlated with overall DOC concentration, and at low DOC levels the relative content of CA was high and vice versa. This followed a seasonal trend, with CA accounting for a smaller proportion of the DOC in winter and a larger part in spring/early summer. A conservative estimate suggested that the studied simple organic carbon compounds potentially could cover 30% of the bacterial gross production in the lake and therefore potentially also was an important source of CO2 that occur in supersaturated concentrations in the lake.
Unproductive lakes are generally supersaturated with carbon dioxide (CO2) and emit CO2 to the atmosphere continuously during ice-free periods. However, temporal variation of the partial pressure of CO2 (pCO2) and thus of CO2 evasion to atmosphere is poorly documented. We therefore carried out temporally high-resolution (every 6 h) measurements of the pCO2 using an automated logger system in the surface water of a subarctic, unproductive, lake in the birch forest belt. The study period was June–September 2004. We found that the pCO2 showed large seasonal variation, but low daily variation. The seasonal variation was likely mainly caused by variations in input and mineralization of allochthonous organic matter. Stratification depth probably also influenced pCO2 of the surface water by controlling the volume in which mineralization of dissolved organic carbon (DOC) occurred. In lakes, with large variations in pCO2, as in our study lake a high (weekly) sampling intensity is recommended for obtaining accurate estimates of the evasion of CO2.
Measurements of the gas transfer rate of CO2 between lake water and the atmosphere present a critical problem for the understanding of lake ecosystem carbon balances and landscape carbon budgets. We present calculations of the gas transfer rate of CO2 from direct measurements of the CO2 flux using an eddy covariance system and concurrent measurements of the concentration of CO2 in the surface water in a lake in boreal zone of northern Sweden. The measured gas transfer rate was different, and in general larger than, rates obtained with the most commonly used models for prediction of the gas transfer rate in lakes. The normalized gas transfer rate (k600EC) was well predicted from the wind speed at 10 m height if data were bin classed into wind classes of 1 m/s for winds above 1 m/s. Unbinned data were also correlated to wind speed but also to water temperature, water temperature/air temperature ratio and to incoming photosynthetic active radiation (PAR). These relationships could reflect effects of both physico-chemical reactions and biological activity.
We quantified the utilization of terrestrial organic matter (OM) in the food web of a humic lake by analyzing the metabolism and the consumers' stable isotopic (C, H, N) composition in benthic and pelagic habitats. Terrestrial OM inputs (3 g C m(-2) d(-1)) to the lake greatly exceeded autochthonous OM production (3 mg C m(-2) d(-1)) in the lake. Heterotrophic bacterial growth (19 mg C m(-2) d(-1)) and community respiration (115 mg C m(-2) d(-1)) were high relative to algal photosynthesis and were predominantly (> 85%) supported by terrestrial OM in both habitats. Consequently, terrestrial OM fueled most (85%) of the total production at the base of the lake's food web (i.e., the sum of primary and bacterial production). Despite the uncertainties of quantitatively estimating resource use based on stable isotopes, terrestrial OM clearly also supported around half the zooplankton (47%), macrozoobenthos (63%), and fish (57%) biomass. These results indicate that, although rates of terrestrial OM inputs were around three orders of magnitude greater than that of autochthonous OM production, the use of the two resources by higher trophic levels was roughly equal. The disproportionally low reliance on terrestrial OM at higher trophic levels, compared with its high rates of input and high support of basic biomass production in the lake, suggests that autochthonous resources could not be completely replaced by terrestrial resources and indicates an upper limit to terrestrial support of lake food webs.
We carried out short-term (2 d) experiments in nine unproductive lakes in northern Sweden in order to investigate organic carbon sources supporting lake water respiration. Surface water was incubated in gas-tight bottles in the dark, and the concentration and isotopic composition (d13C) of dissolved inorganic carbon (DIC) were measured at the start and end of the incubations. Keeling plot analyses revealed that the d13C of the respired carbon was between -28.4% and -30.6% in the lakes and that the respired carbon was mainly of allochthonous organic carbon (AlloOC) origin. The respiration of AlloOC corresponded well with metabolic imbalances indicated by negative net ecosystem production (NEP) values in the lake waters. Keeling plot analysis of DIC accumulating in the hypolimnion of two lakes during summer stratification showed d13C values of around -26.6% for excess DIC, implying that the accumulation of DIC was mainly derived from respiration of AlloOC. Our data provide direct evidence that net heterotrophy of these lakes is caused by input and respiration of AlloOC. We conclude that the Keeling plot method is a powerful technique that enables characterization and quantification of the organic carbon sources contributing to respiration in aquatic systems.
We examined the relationship between planktonic primary production (PP) and bacterial production (BP) in 16 subarctic lakes along an altitude gradient extending from colored coniferous forest lakes to clearwater high alpine lakes. We tested the hypothesis that there was a shift from low to high PP:BP ratios along this gradient. The clearwater alpine lakes had low PP:BP ratios, generally well below 1.0, while the highest ratios were-found in the forest lakes. In contradiction to our hypothesis, the pelagic systems of the clearwater lakes were thus dominated by bacterial energy mobilization from external carbon sources. In this respect the alpine lakes were similar to highly humic lakes. We suggest that the relationship between C, N, and P plays a critical role in determining the PP:BP ratio, and that the N:P ratio in particular can be critical for development of PP or BP dominance in the pelagic systems of unproductive lakes.
We examined changes in bacterioplankton standing stock and production in subarctic lakes in the north of Sweden to elucidate their coupling to lake physical, chemical, and biological characteristics. Sixteen lakes situated along an altitude gradient extending from the coniferous forest to the high-alpine belt were studied during 1998 and 1999. The summer mean bacterial numbers and production varied substantially between the lakes, with a general trend toward decreasing values with increasing altitude. The results demonstrate that P probably restricted bacterial utilization of DOC in the coniferous forest lakes, while low DOC concentrations limited bacterial growth during the summer in the alpine lakes. The primary production of plankton was insufficient to support bacterial production in the lakes. High input of allochthonous DOC to the alpine lakes in spring was sufficient both to increase the bacterial production and to induce P-limitation. As a consequence, there was a tendency toward higher bacterial activity in the spring compared to the summer in the alpine lakes. The results indicate that most of the bacterial standing stock and production are supported by allochthonous DOC plus DOC from benthic production, and more or less limited by the phosphorus supply. We therefore suggest that bacteria populations in subarctic lakes may be indirectly affected by climate variations through its impact on the input of DOC and nutrients from the lake catchments.
Climate change is predicted to be dramatic at high latitudes. Still, climate impact on high latitude lake ecosystems is poorly understood. We studied 15 subarctic lakes located in a climate gradient comprising an air temperature difference of about 6&DEG; C. We show that lake water productivity varied by one order of magnitude along the temperature gradient. This variation was mainly caused by variations in the length of the ice-free period and, more importantly, in the supply of organic carbon and inorganic nutrients, which followed differences in terrestrial vegetation cover along the gradient. The results imply that warming will have rapid effects on the productivity of high latitude lakes, by prolongation of ice-free periods. However, a more pronounced consequence will be a delayed stimulation of the productivity following upon changes of the lakes terrestrial surroundings and subsequent increasing input of elements that stimulate the production of lake biota.
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.
1. To assess the use of stable nitrogen isotopes (δ15N) for reconstructing trophic relationships in planktonic food webs, crustacean zooplankton species and pelagic dissolved and particulate matter were analysed in 14 subarctic lakes in northern Sweden. The lakes are situated along an altitudinal gradient and show a substantial variation in nutrient content and energy mobilization by bacterioplankton and phytoplankton.
2. The δ15N of dissolved and particulate matter was comparatively low, suggesting efficient N recycling and low losses of depleted N from the pelagic zone of these unproductive lakes.
3. Copepods had a systematically higher δ15N than cladocerans, with an average difference of 3.1–4.9‰ within lakes, implying different trophic positions of the two groups. Comparisons of nitrogen pools and energy fluxes suggest that the low cladoceran δ15N was a result of feeding on bacteria.
4. The difference in δ15N between copepods and cladocerans declined with decreasing bacterioplankton production among lakes, due either to increasing trophic isotope fractionation or decreasing relative importance of bacteria in the diet of cladocerans.
Global warming and land use scenarios suggest increased 21st 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.
Forestry practices often result in an increased export of carbon and nitrogen to downstream aquatic systems. Although these losses affect the greenhouse gas (GHG) budget of managed forests, it is unknown if they modify GHG emissions of recipient aquatic systems. To assess this question, air-water fluxes of carbon dioxide (CO2), methane (CH4) and nitrous oxide (N2O) were quantified for humic lakes and their inlet streams in four boreal catchments using a before-after control-impact experiment. Two catchments were treated with forest clear-cuts followed by site preparation (18 % and 44 % of the catchment area). GHG fluxes and hydrological and physicochemical water characteristics were measured at multiple locations in lakes and streams at high temporal resolution throughout the summer season over a 4-year period. Both lakes and streams evaded all GHGs. The treatment did not significantly change GHG fluxes in streams or lakes within 3 years after the treatment, despite significant increases of CO2 and CH4 concentrations in hillslope groundwater. Our results highlight that GHGs leaching from forest clear-cuts may be buffered in the riparian zone-stream continuum, likely acting as effective biogeochemical processors and wind shelters to prevent additional GHG evasion via downstream inland waters. These findings are representative of low productive forests located in relatively flat landscapes where forestry practices cause only a limited initial impact on catchment hydrology and biogeochemistry.
Natural organic matter (NOM) from stream and soil water in a humic-rich headwater catchment in northern Sweden (initial total organic carbon (TOC) concentrations 10-40 mg C L-1) was rapidly degraded by light and microbial activity in an incubation experiment. Concentration losses were 33-50% after 12 days of exposure to 69 W m(-2) artificial PAR and 16 W m(-2) UV radiation. Natural, unshaded mid-day solar radiation in the region (68degreesN 18degreesE) during the month of june is 159 W m(-2) for PAR. In contrast to microbial organic carbon removal, TOC exponentially decreased upon radiation, which suggests that TOC is more rapidly oxidized by light than by ambient microbes. Further, rapid decline in TOC concentration implies the presence of a dominant pool of photo-labile compounds (p > 95%). A measured mass balance for carbon identified 50-75% of the degraded TOC as carbon dioxide after 12 days of exposure to light. The observed conversion of organic to inorganic carbon was accompanied by increases in pH and alkalinity, suggesting that photo-degradation of NOM potentially contributes to in-stream buffering capacity. The remaining refractory TOC changed in chemical character, including an altered molecular weight distribution with decreased average weight and a change in the proportions of humics as evidenced by absorbance ratios (A(254)/A(420)). Extrapolation of the experiment to natural headwater conditions show that photo-degradation is an important in-stream process that should be considered in calculations of carbon turnover in surface waters because of its influence on both TOC amount and character.
Global environmental changes are causing widespread nutrient depletion, declines in the ratio of dissolved inorganic nitrogen (N) to total phosphorus (DIN:TP), and increases in both water temperature and terrestrial colored dissolved organic carbon (DOC) concentration (browning) in high-latitude northern lakes. Declining lake DIN:TP, warming, and browning alter the nutrient limitation regime and biomass of phytoplankton, but how these stressors together affect the nutritional quality in terms of polyunsaturated fatty acid (PUFA) contents of the pelagic food web components remains unknown. We assessed the fatty acid compositions of seston and zooplankton in 33 lakes across south-to-north and boreal-to-subarctic gradients in Sweden. Data showed higher lake DIN:TP in the south than in the north, and that boreal lakes were warmer and browner than subarctic lakes. Lake DIN:TP strongly affected the PUFA contents—especially eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA)—in seston, calanoids, and copepods (as a group), but not in cladocerans. The EPA+DHA contents increased by 123% in seston, 197% in calanoids, and 230% in copepods across a lake molar DIN:TP gradient from 0.17 to 14.53, indicating lower seston and copepod nutritional quality in the more N-limited lakes (those with lower DIN:TP). Water temperature affected EPA+DHA contents of zooplankton, especially cladocerans, but not seston. Cladoceran EPA+DHA contents were reduced by ca. 6% for every 1°C increase in surface water. Also, the EPA, DHA, or EPA+DHA contents of Bosmina, cyclopoids, and copepods increased in lakes with higher DOC concentrations or aromaticity. Our findings indicate that zooplankton food quality for higher consumers will decrease with warming alone (for cladocerans) or in combination with declining lake DIN:TP (for copepods), but impacts of these stressors are moderated by lake browning. Global environmental changes that drive northern lakes toward more N-limited, warmer, and browner conditions will reduce PUFA availability and nutritional quality of the pelagic food web components.
Gas fluxes from lakes and other stratified water bodies, computed using conservative values of the gas transfer coefficient k600, have been shown to be a significant component of the carbon cycle. We present a mechanistic analysis of the dominant physical processes modifying k600 in a stratified lake and resulting new models of k600 whose use will enable improved computation of carbon fluxes. Using eddy covariance results, we demonstrate that i) higher values of k600 occur during low to moderate winds with surface cooling than with surface heating; ii) under overnight low wind conditions k600 depends on buoyancy flux β rather than wind speed; iii) the meteorological conditions at the time of measurement and the inertia within the lake determine k600; and iv) eddy covariance estimates of k600 compare well with predictions of k600 using a surface renewal model based on wind speed and β.
Anthropogenic environmental stressors (like atmospheric deposition, land use change, and climate warming) are predicted to increase inorganic nitrogen and organic carbon loading to northern boreal lakes, with potential consequences for denitrification in lakes. However, our ability to predict effects of these changes is currently limited as northern boreal lakes have been largely neglected in denitrification studies. The aim of this study was therefore to assess how maximum potential denitrification and N2O production rates, and the relationship between the two (relative N2O production), is controlled by availability of nitrate (NO3−), carbon (C), phosphorus (P), and temperature. Experiments were performed using the acetylene inhibition technique on sediments from a small, nutrient poor boreal lake in northern Sweden in 2014. Maximum potential denitrification and N2O production rates at 4°C were reached already at NO3− additions of 106–120 μg NO3−–N/L, and remained unchanged with higher NO3 amendments. Higher incubation temperatures increased maximum potential denitrification and N2O production rates, and Q10 was somewhat higher for N2O production (1.77) than for denitrification (1.69). The relative N2O production ranged between 13% and 64%, and was not related to NO3− concentration, but the ratio increased when incubations were amended with C and P (from a median of 16% to 27%). Combined, our results suggests that unproductive northern boreal lakes currently have low potential for denitrification but are susceptible to small changes in NO3 loading especially if these are accompanied by enhanced C and P availability, likely promoting higher N2O production relative to N2.
The organic carbon balance of a lake with high input of allochthonous organic carbon is modelled integrating physical, chemical and biological processes. The physical model captures the behaviour of real thermal stratification in the lake for different flow situations during the period 1993-1997. The dissolved organic carbon model is based on simulated trajectories of water parcels. By tracking parcels, account is kept of environmental factors such as temperature and radiation as well as DOC quality for each parcel, The DOC concentration shows seasonal variations primarily dependent on inflow. The organic matter degradation (bacterial- and photodegradation) in the lake amounts to 1.5-2.5 mg C l(-1) yr(-1), where photooxidation is responsible for approximately 10%. The estimated DIC production in the lake is large compared to sediment mineralisation and primary production. The main conclusion is that the model with the selected parameterisations of the degradation processes reasonably well describes the DOC dynamics in a forest lake.
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.
CO2 evasion from freshwater lakes is an important component of the carbon cycle. However, the relative contribution from different lake sizes may vary, since several parameters underlying CO2 flux are size dependent. Here we estimated the annual lake CO2 evasion from a catchment in northern Sweden encompassing about 30,000 differently sized lakes. We show that areal CO2 fluxes decreased rapidly with lake size, but this was counteracted by the greater overall coverage of larger lakes. As a result, total efflux increased with lake size and the single largest lake in the catchment dominated the CO2 evasion (53% of all CO2 evaded). By contrast, the contribution from the smallest ponds (about 27,000) was minor (<6%). Our results emphasize the importance of accounting for both CO2 flux rates and areal contribution of various sized lakes in assessments of CO2 evasion at the landscape scale.
Variation of the surface water CO2 concentration is likely to be the result of biological activity and physical processes as water mixing and gas exchange with the atmosphere. Here we have studied the variations in surface water CO2 during the ice-free period in the humic Lake Merasjärvi in northern Sweden. Meteorological, hydrological and limnological data were collected using data logging equipment permitting high time-resolution. The surface water of the lake was supersaturated with respect to CO2 throughout the study period. There were, however, considerable diurnal and longer-term temporal variations of the surface water CO2 concentrations. Partial least squares (PLS) models were used to link the logged CO2 data to the multivariate dataset. On the longer-term time scale (analyzed with 24h means of the logged data) high concentrations of surface water CO2 were best related to the depth and temperature of the upper warmer layer (epilimnion), and to erosion of the underlying colder layer (hypolimnion). The diurnal variation (analyzed with 30 minute means of the logged data) was best related to the thermal dynamics within the epilimnion, which regulated the surface water access to CO2 stores within this layer. Variables related to CO2 emission and photosynthesis (wind and PAR), showed only weak correlations to variations of the surface water CO2 concentration. Accordingly, the CO2 flux, measured with the eddy-covariance technique, was not correlated to the surface water CO2 concentration.
1. Both the pelagic and benthic net dissolved inorganic carbon (DIC) productions were measured in situ on four occasions from June to September 2004, in the unproductive Lake Diktar-Erik in subarctic Sweden. The stable isotopic signal (δ13C) of respired organic material was estimated from hypolimnion water data and data from a laboratory incubation using epilimnion water.
2. Both pelagic and benthic habitats were net heterotrophic during the study period, with a total net DIC production of 416 mg C m-2 day-1, of which the pelagic habitat contributed approximately 85%. The net DIC production decreased with depth both in the pelagic water and in the sediments, and most of the net DIC production occurred in the upper water column.
3. Temporal variations in both pelagic and benthic DIC production were small, although we observed a significant decrease in pelagic net DIC production after the autumn turnover. Water temperature was the single most important factor explaining temporal and vertical variations in pelagic DIC production. No single factor explained more than 10% of the benthic net DIC production, which probably was regulated by several interacting factors.
4. Pelagic DIC production, and thus most of the whole-lake net production of DIC, was mainly due to the respiration of allochthonous organic carbon. Stable isotope data inferred that nearly 100% of accumulated DIC in the hypolimnion water had an allochthonous carbon source. Similarly, in the laboratory incubation using epilimnion water, c. 85% of accumulated DIC was indicated to have an allochthonous organic carbon source.