Lake restoration influences nutritional quality of algae and consequently Daphnia biomass

AbstractFood quality is one of the key factors influencing zooplankton population dynamics. Eutrophication drives phytoplankton communities toward the dominance of cyanobacteria, which means a decrease in the availability of sterols and long-chain polyunsaturated fatty acids (EPA and DHA). The effects of different restoration measures on the nutritional quality of the phytoplankton community and subsequent impacts on zooplankton biomass have rarely been considered. We analyzed the nutritional quality of phytoplankton in the eutrophic Lake Vesijärvi in southern Finland over a 37-year period, and studied the impacts of two restoration measures, biomanipulation and hypolimnetic aeration, on th…

Terrestrial organic matter quantity or decomposition state does not compensate for its poor nutritional quality for Daphnia

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…

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…

Retroconversion of docosapentaenoic acid (n-6): an alternative pathway for biosynthesis of arachidonic acid in Daphnia magna.

The aim of this study was to assess metabolic pathways for arachidonic acid (20:4n-6) biosynthesis in Daphnia magna. Neonates of D. magna were maintained on [13C] enriched Scenedesmus obliquus and supplemented with liposomes that contained separate treatments of unlabeled docosapentaenoic acid (22:5n-6), 20:4n-6, linoleic acid (18:2n-6) or oleic acid (18:1n-9). Daphnia in the control treatment, without any supplementary fatty acids (FA) containing only trace amounts of 20:4n-6 (~0.3 % of all FA). As expected, the highest proportion of 20:4n-6 (~6.3 %) was detected in Daphnia that received liposomes supplemented with this FA. Higher availability of 18:2n-6 in the diet increased the proportio…

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…

A Fatty Acid Based Bayesian Approach for Inferring Diet in Aquatic Consumers

We modified the stable isotope mixing model MixSIR to infer primary producer contributions to consumer diets based on their fatty acid composition. To parameterize the algorithm, we generated a 'consumer-resource library' of FA signatures of Daphnia fed different algal diets, using 34 feeding trials representing diverse phytoplankton lineages. This library corresponds to the resource or producer file in classic Bayesian mixing models such as MixSIR or SIAR. Because this library is based on the FA profiles of zooplankton consuming known diets, and not the FA profiles of algae directly, trophic modification of consumer lipids is directly accounted for. To test the model, we simulated hypothet…

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…

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…

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

Article

Combined effects of eutrophication and warming on polyunsaturated fatty acids in complex phytoplankton communities:A mesocosm experiment

Climate change and eutrophication are among the main stressors of shallow freshwater ecosystems, and their effects on phytoplankton community structure and primary production have been studied extensively. However, their combined effects on the algal production of polyunsaturated fatty acids (PUFA), specifically, eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) are currently unresolved. Moreover, the proximate reasons for changes in phytoplankton EPA and DHA concentrations are unclear, i.e., the relative importance of ecological (changes in the community composition) vs. ecophysiological (within taxa changes in EPA and DHA levels) factors. We investigated the responses of phytopla…

Terrestrial carbohydrates support freshwater zooplankton during phytoplankton deficiency

Article

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…

Fatty acid composition as biomarkers of freshwater microalgae: analysis of 37 strains of microalgae in 22 genera and in seven classes

The fatty acid (FA) composition of algae is an important determinant of their food quality for consumers, and FAs can also be used as biomarkers for biochemical and energetic pathways in food webs. FA analyses of 7 freshwater algal classes and 37 strains showed clear similarity within classes and strong differences amongst classes. Class was a dominant factor (66.4%) explaining variation in FA signatures of microalgae. The 7 algal classes comprised 4 separate groups according to their FA profiles: (1) Chlorophyceae and Trebouxiophyceae, (2) Bacillariophyceae, (3) Cryptophyceae, Chrysophyceae, and Raphidophyceae, and (4) Euglenophyceae. Each group had a characteristic FA composition, althoug…

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 …

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…

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.

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.