Oxidation, efflux, and isotopic fractionation of methane during autumnal turnover in a polyhumic, boreal lake

[1] We studied the oxidation and efflux of methane (CH4) in a small, polyhumic lake, Mekkojarvi (southern Finland), during 6 weeks in autumn when the stability of the water mass first weakened, temporarily restabilized, and finally mixed completely. During the summer stratification period, CH4 had accumulated in the anoxic hypolimnion to high concentrations (>150 mmol m−3). Gradual mixing of the water column during the autumn allowed access to both oxygen and CH4 by aerobic methane-oxidizing bacteria (MOB) deeper in the water column. Thus the bulk (∼83–88%) of the CH4 accumulated in the hypolimnion was subsequently consumed by MOB while only 12–17% was lost from the lake to the atmosphere a…

Recycling and fluxes of carbon gases in a stratified boreal lake following experimental carbon addition

Partly anoxic stratified humic lakes are important sources of methane (CH4) and carbon dioxide (CO2) to the atmosphere. We followed the fate of CH4 and CO2 in a small boreal stratified lake, Alinen Mustajärvi, during 2007–2009. In 2008 and 2009 the lake received additions of dissolved organic carbon (DOC) with stable carbon isotope ratio (δ13C) around 16‰ higher than that of local allochthonous DOC. Carbon transformations in the water column were studied by measurements of δ13C of CH4 and of the dissolved inorganic carbon (DIC). Furthermore, CH4 and CO2 production, consumption and emissions were estimated. Methane oxidation was estimated by a diffusion gradient method. The amount, lo…

Whole-lake dissolved inorganic 13C additions reveal seasonal shifts in zooplankton diet.

Sustained whole-lake additions of 13C-enriched dissolved inorganic carbon (DIC), intended to increase experimentally the delta13C of DIC in the epilimnion of a small lake with high dissolved organic carbon (DOC), were made during three seasonal periods (spring, summer, and autumn). Coupled with carbon and nitrogen stable isotope analysis of zooplankton and several of their putative food sources, these additions were used to investigate seasonal changes in the relative contributions of different food sources to zooplankton diet in the lake. Four main potential food sources were considered: phytoplankton, heterotrophic bacteria (HB), methanotrophic bacteria (MOB), and green sulfur bacteria (G…

Seasonal shifts in the diet of lake zooplankton revealed by phospholipid fatty acid analysis

SUMMARY 1. We analysed the phospholipid fatty acid (PLFA) profiles of seston and of the dominant zooplankter, Daphnia longispina, through the open water period in a small, dystrophic lake to investigate seasonal variation in the diet of Daphnia. Phytoplankton, heterotrophic bacteria, green sulphur bacteria and methane-oxidizing bacteria (MOB) were all present in the water column of the lake, and previous studies have indicated that vertically migrating Daphnia can exploit all these potential food sources. 2. For adult Daphnia, although there was some correspondence between the PLFA profile of Daphnia and the concurrent seston PLFA profile, strongest correlations were between the Daphnia PLF…

Lake browning impacts community structure and essential fatty acid content of littoral invertebrates in boreal lakes

AbstractMany lakes in the northern hemisphere are browning due to increasing concentrations of terrestrial dissolved organic carbon (DOC). The consequences of lake browning to littoral invertebrates, however, are not fully understood. We analyzed community structure and fatty acid (FA) profiles of littoral invertebrates in humic (DOC-rich) and clear-water lakes in Eastern Finland. We found higher abundance of chironomids (Diptera: Chironomidae) in humic compared to clear-water lakes, whereas stoneflies (Plecoptera) and mayflies (Ephemeroptera: Baetidae) were more abundant in clear-water lakes. Taxon explained 65% of the differences in the FA composition of littoral invertebrates. However, t…

Methane-oxidizing and photoautotrophic bacteria are major producers in a humic lake with large anoxic hypolimnion

Heterotrophic processes are considered to prevail in lakes with high terrestrial loading and a high content of dissolved organic carbon. However, previous studies have shown abundant methanotrophic bacteria (MOB) and photoautotrophic green sulphur bacteria (GSB) in some boreal lakes. We used genetic and lipid biomarkers to follow the seasonal succession of the abundance and carbon biomass of the major bacterial groups in Lake Mekkojarvi, a small Finnish polyhumic lake with a large anoxic hypolimnion. Biomarkers were based on length heterogeneity analysis of 16S rRNA gene amplification by polymerase chain reaction (PCR) (LH-PCR) and on phospholipid fatty acid (PLFA) analysis. Representative …

Nutritional quality of littoral macroinvertebrates and pelagic zooplankton in subarctic lakes

Littoral benthic primary production is considered the most important energy source of consumers in subarctic lakes. We analyzed essential fatty acid (EFA) and amino acid (EAA) content of 23 littoral benthic macroinvertebrate taxa as well as cladocerans and copepods from pelagic and littoral habitats of 8–9 subarctic lakes to compare their nutritional quality. Pelagic crustacean zooplankton had significantly higher EFA and total FA content (on average 2.6‐fold and 1.6‐fold, respectively) than littoral macroinvertebrates in all our study lakes. Specifically, docosahexaenoic acid (DHA), one of the most important EFA for juvenile fish, was almost exclusively found in pelagic copepods. In littor…

Epizooic ciliates (Vorticella sp.) compete for food with their host Daphnia longispina in a small polyhumic lake.

Clearance rates of epizooic ciliates (Vorticella sp.) were measured together with their host, a planktonic cladoceran Daphnia longispina by using fluorescent latex beads as tracers of food. Vorticellans and their host graze on food of same size range (nanoplanktonic algae and bacteria). Individual clearance rates of Vorticella averaged 6.9 and 7.0 μl ind-1 h-1 and those of Daphnia 463 and 708 μl ind-1 h-1 for beads with diameter 2.00 and 3.92 μm. On the average, epizooic vorticellans together on the carapace of Daphnia cleared particles with rates representing 25–33% of that the host cleared, the maximum rates being 50–80%. In a steeply stratified polyhumic lake vorticellans take advantage …

Land Use Affects Carbon Sources to the Pelagic Food Web in a Small Boreal Lake

Small humic forest lakes often have high contributions of methane-derived carbon in their food webs but little is known about the temporal stability of this carbon pathway and how it responds to environmental changes on longer time scales. We reconstructed past variations in the contribution of methanogenic carbon in the pelagic food web of a small boreal lake in Finland by analyzing the stable carbon isotopic composition (δ13C values) of chitinous fossils of planktivorous invertebrates in sediments from the lake. The δ13C values of zooplankton remains show several marked shifts (approx. 10 ‰), consistent with changes in the proportional contribution of carbon from methane-oxidizing bacteri…

Effects of alternative electron acceptors on the activity and community structure of methane-producing and consuming microbes in the sediments of two shallow boreal lakes

The role of anaerobic CH4 oxidation in controlling lake sediment CH4 emissions remains unclear. Therefore, we tested how relevant EAs (SO42−, NO3−, Fe3+, Mn4+, O2) affect CH4 production and oxidation in the sediments of two shallow boreal lakes. The changes induced to microbial communities by the addition of Fe3+ and Mn4+ were studied using next-generation sequencing targeting the 16S rRNA and methyl-coenzyme M reductase (mcrA) genes and mcrA transcripts. Putative anaerobic CH4-oxidizing archaea (ANME-2D) and bacteria (NC 10) were scarce (up to 3.4% and 0.5% of archaeal and bacterial 16S rRNA genes, respectively), likely due to the low environmental stability associated with shallow depths.…

Population structure, life cycle, and trophic niche of the glacial relict amphipod, Gammaracanthus lacustris, in a large boreal lake

Ecology of the glacial relict macrocrustacean Gammaracanthus lacustris, a rare inhabitant of deep Fennoscandian lakes, is poorly understood. We studied the life cycle and trophic position of this cold‐stenothermic amphipod in Lake Paasivesi, eastern Finland. The study is based on intensive sampling and analyses of fatty acid composition as well as stable carbon (δ13C) and nitrogen (δ15N) isotope ratios. Both day and night, the G. lacustris population occurred at depths below 25 m at temperatures 40 mm and live up to 4 years. The oldest and the largest individuals and females seemed to favour the deepest zones. In October, almost 100% of females with length at least 25 mm (i.e. females presu…

Diet-specific biomarkers show that high-quality phytoplankton fuels herbivorous zooplankton in large boreal lakes

SUMMARY 1. The zooplankton is a key link in the transfer of energy from primary producers up through aquatic food webs. Previous efforts to quantify the importance of basal resources to aquatic consumers have used stable isotopes (SI) and simple ternary models, including only ‘bulk’ phytoplankton, bacteria or terrestrial particulate organic matter (t-POM). 2. We used a novel Bayesian mixing model based on fatty acids (FA) to quantify the dietary assimilation of seven basal resources, including five phytoplankton groups, pelagic bacteria and t-POM, to Cladocera in large boreal lakes in Finland. To account for trophic enrichment of FA from the diet to consumers, we parameterised the model wit…

Whole-Lake Sugar Addition Demonstrates Trophic Transfer of Dissolved Organic Carbon to Top Consumers

Terrestrial dissolved organic carbon (DOC) provides an external carbon source to lake ecosystems. However, there is ongoing debate about whether external DOC that enters a lake can pass up the food web to support top consumers. We show, from experimental manipulation of a whole lake, that externally loaded DOC can contribute appreciably to fish biomass. Monthly additions of cane sugar with a distinct carbon stable isotope value during 2 years rapidly enriched the 13C content of zooplankton and macroinvertebrates, with a more gradual 13C enrichment of fish. After sugar addition stopped, the 13C content of consumers reverted towards original values. A simple isotope mixing model indicated tha…

Population structure, life cycle, and trophic niche of the glacial relict amphipod, Gammaracanthus lacustris , in a large boreal lake

Taxonomic identity and phytoplankton diet affect fatty acid composition of zooplankton in large lakes with differing dissolved organic carbon concentration

An increase in loading of dissolved organic carbon (DOC) from terrestrial to freshwater ecosystems has recently been documented in several boreal regions. We studied how transfer of important biomolecules, fatty acids, may be connected to DOC in pelagic food webs of large boreal lakes. We collected nine zooplankton taxa during three seasons from six lakes along a gradient of DOC (5–10 mg C L−1). Taxonomic identity explained 62% of the variation in zooplankton fatty acids, while the effects of DOC and season were less pronounced. This suggests that the community composition of zooplankton is important in determining the quality of food available for higher trophic level consumers. Seasonal d…

Periphyton as a key diet source of essential fatty acids for macroinvertebrates across a nutrient and dissolved organic carbon gradient in boreal lakes

We studied how physiologically important long-chain polyunsaturated fatty acids (PUFA) in benthic macroinvertebrates (Asellus aquaticus, Chironomidae, and Oligochaeta) were related to those in periphyton and terrestrial organic matter (tree leaves), collected from littoral areas of 17 boreal lakes that differed in their dissolved organic carbon (DOC) and nutrient (phosphorus and nitrogen) concentrations. We also analyzed fatty acid (FA)-specific stable carbon isotopes (δ13CFA) to investigate the dietary origin (periphyton vs. terrestrial organic matter) of PUFA in the consumers. In contrast to periphyton, terrestrial organic matter was deprived of long-chain PUFA, such as eicosapentaenoic a…

Planktivorous vendace (Coregonus albula) utilise algae-derived fatty acids for biomass increase and lipid deposition

Spatial and seasonal variation in the abundance and composition of phytoplankton and other basal resources (bacteria, terrestrial detritus) influence the availability of essen-tial polyunsaturated fatty acids (PUFA) for upper trophic levels in lake food webs. We studied accumulation, composition and content of fatty acids in planktivorous ven-dace (Coregonus albula) muscle tissue during the summer feeding period (May to late September) in six boreal lakes. Vendace muscle fatty acid content increased from ~17.7 mg/g DW to ~28.1 mg/g DW from spring to summer, corresponding to a ~59% increase in total fatty acids. PUFA accounted for 45%–65% of all fatty acids, and the most abundant fatty acid …

Browning-induced changes in trophic functioning of planktonic food webs in temperate and boreal lakes: insights from fatty acids

AbstractThe effects of lake browning on trophic functioning of planktonic food webs are not fully understood. We studied the effects of browning on the response patterns of polyunsaturated fatty acids and n−3/n−6 ratio in seston and compared them between boreal and temperate lakes. We also compared the regional differences and the effects of lake browning on the reliance of zooplankton on heterotrophic microbial pathways and the mass fractions of eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) in zooplankton. Lake browning was associated with increasing phytoplankton biomass and concentrations of EPA and DHA in both temperate and boreal lakes, but the seston n−3/n−6 ratio was low…

Experimental δ13C evidence for a contribution of methane to pelagic food webs in lakes

We tested the hypothesis that low stable carbon isotope (δ13C) values commonly observed for zooplankton in humic lakes are due to their feeding on isotopically light methane-oxidizing microbes, and thus that methane-derived carbon is important in the food webs of these lakes. In replicate laboratory cultures, Daphnia longispina, a common crustacean zooplankter in humic lakes, were fed microbial suspensions with or without enrichment by biogenic methane. The δ13C values of Daphnia indicated consumption of 13C-depleted methanotrophic bacteria, while growth rates, survival, and reproduction of Daphnia in cultures enriched with methane were equal to or greater than those in nonenriched cultures…

DifferingDaphnia magnaassimilation efficiencies for terrestrial, bacterial, and algal carbon and fatty acids

There is considerable interest in the pathways by which carbon and growth-limiting elemental and biochemical nutrients are supplied to upper trophic levels. Fatty acids and sterols are among the most important molecules transferred across the plant-animal interface of food webs. In lake ecosystems, in addition to phytoplankton, bacteria and terrestrial organic matter are potential trophic resources for zooplankton, especially in those receiving high terrestrial organic matter inputs. We therefore tested carbon, nitrogen, and fatty acid assimilation by the crustacean Daphnia magna when consuming these resources. We fed Daphnia with monospecific diets of high-quality (Cryptomonas marssonii) a…

Increasing concentration of polyunsaturated fatty acids in browning boreal lakes is driven by nuisance algaGonyostomum

Elevated concentrations of dissolved organic carbon (DOC) promote browning of lakes, which alters the physicochemical properties of water and ecosystem functioning. However, browning‐induced changes in basal production of polyunsaturated fatty acids from the n‐3 and n‐6 families (n‐3 and n‐6 PUFA) in lakes are not fully understood. The concentrations of PUFA, which are micronutrients required to maintain growth and reproduction of consumers, have been suggested to either rise or decline in seston as a response to lake browning. Elevated DOC concentrations may also promote bacterial biomass and production and thus potentially increase the concentration of bacterial fatty acids (BAFA) in sest…

Lake eutrophication and brownification downgrade availability and transfer of essential fatty acids for human consumption

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How important are terrestrial organic carbon inputs for secondary production in freshwater ecosystems?

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Terrestrial carbohydrates support freshwater zooplankton during phytoplankton deficiency

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Differing Daphnia magna assimilation efficiencies for terrestrial, bacterial, and algal carbon and fatty acids

There is considerable interest in the pathways by which carbon and growthlimiting elemental and biochemical nutrients are supplied to upper trophic levels. Fatty acids and sterols are among the most important molecules transferred across the plant–animal interface of food webs. In lake ecosystems, in addition to phytoplankton, bacteria and terrestrial organic matter are potential trophic resources for zooplankton, especially in those receiving high terrestrial organic matter inputs. We therefore tested carbon, nitrogen, and fatty acid assimilation by the crustacean Daphnia magna when consuming these resources. We fed Daphnia with monospecific diets of high-quality (Cryptomonas marssonii) an…

Inferring phytoplankton community composition with a fatty acid mixing model

The taxon specificity of fatty acid composition in algal classes suggests that fatty acids could be used as chemotaxonomic markers for phytoplankton composition. The applicability of phospholipid-derived fatty acids as chemotaxonomic markers for phytoplankton composition was evaluated by using a Bayesian fatty acid-based mixing model. Fatty acid profiles from monocultures of chlorophytes, cyanobacteria, diatoms, euglenoids, dinoflagellates, raphidophyte, cryptophytes and chrysophytes were used as a reference library to infer phytoplankton community composition in five moderately humic, large boreal lakes in three different seasons (spring, summer and fall). The phytoplankton community compo…

Trophic upgrading via the microbial food web may link terrestrial dissolved organic matter to Daphnia

Direct consumption of allochthonous resources generally yields poor growth and reproduction in zooplankton, but it is still unclear how trophic upgrading of terrestrial dissolved organic matter (tDOM) via the microbial food web may support zooplankton. We compared survival, somatic growth and reproduction of Daphnia magna fed with the heterotrophic flagellate Paraphysomonas vestita and three algal diets. Paraphysomonas was fed lake bacteria that used tDOM as a substrate to simulate an allochthonous diet that zooplankton encounter in lakes. The highest survival, growth and reproduction was achieved with a diet of Cryptomonas, while Daphnia performance was the worst when fed Microcystis. Para…

Lake zooplankton delta13C values are strongly correlated with the delta13C values of distinct phytoplankton taxa

Article

Allochthony, fatty acid and mercury trends in muscle of Eurasian perch (Perca fluviatilis) along boreal environmental gradients

Environmental change, including joint effects of increasing dissolved organic carbon (DOC) and total phosphorus (TP) in boreal northern lakes could potentially affects food web energy sources and the biochemical composition of organisms. These environmental stressors are enhanced by anthropogenic land-use and can decrease the quality of polyunsaturated fatty acids (PUFAs) in seston and zooplankton, and therefore, possibly cascading up to fish. In contrast, the content of mercury in fish increases with lake browning potentially amplified by intensive forestry practises. However, there is little evidence on how these environmental stressors simultaneously impact beneficial omega-3 fatty acid …

Increasing temperature and productivity change biomass, trophic pyramids and community‐level omega‐3 fatty acid content in subarctic lake food webs

Climate change in the Arctic is outpacing the global average and land-use is intensifying due to exploitation of previously inaccessible or unprofitable natural resources. A comprehensive understanding of how the joint effects of changing climate and productivity modify lake food web structure, biomass, trophic pyramid shape and abundance of physiologically essential biomolecules (omega-3 fatty acids) in the biotic community is lacking. We conducted a space-for-time study in 20 subarctic lakes spanning a climatic (+3.2 degrees C and precipitation: +30%) and chemical (dissolved organic carbon: +10 mg/L, total phosphorus: +45 mu g/L and total nitrogen: +1,000 mu g/L) gradient to test how temp…

Planktivorous vendace (Coregonus albula ) utilise algae-derived fatty acids for biomass increase and lipid deposition

Spatial and seasonal variation in the abundance and composition of phytoplankton and other basal resources (bacteria, terrestrial detritus) influence the availability of essential polyunsaturated fatty acids (PUFA) for upper trophic levels in lake food webs. We studied accumulation, composition and content of fatty acids in planktivorous vendace (Coregonus albula) muscle tissue during the summer feeding period (May to late September) in six boreal lakes. Vendace muscle fatty acid content increased from ~17.7 mg/g DW to ~28.1 mg/g DW from spring to summer, corresponding to a ~59% increase in total fatty acids. PUFA accounted for 45%–65% of all fatty acids, and the most abundant fatty acid wa…

Ecosystem responses to increased organic carbon concentration: comparing results based on long-term monitoring and whole-lake experimentation

Recent increases in terrestrial dissolved organic carbon (DOC) concentrations in northern inland waters have many ecological consequences. We examined available data on carbon cycles and food webs of 2 boreal headwater lakes in southern Finland. Basic limnology and catchment characteristics of a pristine lake, Valkea-Kotinen (VK), were monitored over the past 25 years while the lake has undergone browning and DOC increased from similar to 11 to 13 mg L-1. Pronounced changes in the early 2000s represent a regime shift in DOC concentration and color. Lake Alinen Mustajarvi (AM) was manipulated for 2 years by additions of labile DOC (cane sugar), raising the DOC concentration from similar to 1…

Appendix C. Results of Cox regression survival analysis of experiments.

Results of Cox regression survival analysis of experiments.

Appendix E. Principal component analysis of diets and Daphnia.

Principal component analysis of diets and Daphnia.

Ecosystem responses to increased organic carbon concentration: comparing results based on long-term monitoring and whole-lake experimentation

Recent increases in terrestrial dissolved organic carbon (DOC) concentrations in northern inland waters have many ecological consequences. We examined available data on carbon cycles and food webs of 2 boreal headwater lakes in southern Finland. Basic limnology and catchment characteristics of a pristine lake, Valkea-Kotinen (VK), were monitored over the past 25 years while the lake has undergone browning and DOC increased from ∼11 to 13 mg L−1. Pronounced changes in the early 2000s represent a regime shift in DOC concentration and color. Lake Alinen Mustajärvi (AM) was manipulated for 2 years by additions of labile DOC (cane sugar), raising the DOC concentration from ∼10 to 12 mg L−1, but …

Appendix A. Detailed concentration information of added nutrients in batch and life table experiments.

Detailed concentration information of added nutrients in batch and life table experiments.

Appendix B. Fatty acid and sterol profiles of terrestrial particulate organic carbon (t-POC), bacteria, and phytoplankton diets.

Fatty acid and sterol profiles of terrestrial particulate organic carbon (t-POC), bacteria, and phytoplankton diets.

Appendix D. Correlations between different nutrients of diets and offspring and body size of Daphnia.

Correlations between different nutrients of diets and offspring and body size of Daphnia.

Appendix F. The parameter estimates of t-POC diets for power functions.

The parameter estimates of t-POC diets for power functions.