151. The prevailing view that many humic lakes are net heterotrophic is commonly based on pelagic measurements alone. Poor light conditions in humic lakes are assumed to constrain littoral primary production (PP), such that the littoral zone has been considered an insignificant contributor to whole-lake PP. However, that assumption is based on models and inferences from pelagic processes which do not take littoral zone structure into account. 20Many lakes have an extensive ring of aquatic vegetation lying near the water surface, which provides substratum for epiphytic algae under well-illuminated conditions. 2. We measured both pelagic and littoral PP and community respiration (CR) in Mekkojärvi, a small, highly humic headwater lake, in southern Finland throughout the open water season in 2012. We used a 14 C incorporation technique to measure pelagic PP, while littoral PP was 25 determined using changes in dissolved inorganic carbon (DIC) concentrations during in situ incubations. We then estimated whole-lake PP and CR for both the littoral and pelagic zones.3. We found that littoral PP usually dominated whole-lake PP, contributing >90% to total PP in summer. A mean pelagic production to respiration ratio (GPP:CR) of 0.4 clearly indicated strong net heterotrophy, but a value of 2.2 for the littoral zone indicated strong autotrophy. 30For both habitats combined, the mean whole lake GPP:CR was 1.6, indicating net autotrophy.4. We suggest that littoral PP can contribute significantly to whole-lake PP even in highly humic lakes, and that the littoral contribution can shift some lakes which have been considered net heterotrophic to metabolic balance or even net autotrophy during the ice-free period. Small humic lakes like Mekkojärvi with rings of littoral vegetation are widespread, 35 especially in the boreal zone, and at least for similar lakes, evaluating lake metabolism from pelagic measurements alone is likely to be misleading.3

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°C and precipitation: +30%) and chemical (dissolved organic carbon: +10 mg/L, total phosphorus: +45 µg/L and total nitrogen: +1,000 µg/L) gradient to test how temperature and productivity jointly affect the structure, biomass and community fatty acid content (eicosapentaenoic acid [EPA] and docosahexaenoic acid [DHA]) of whole food webs. Increasing temperature and productivity shifted lake communities towards dominance of warmer, murky‐water‐adapted taxa, with a general increase in the biomass of primary producers, and secondary and tertiary consumers, while primary invertebrate consumers did not show equally clear trends. This process altered various trophic pyramid structures towards an hour glass shape in the warmest and most productive lakes. Increasing temperature and productivity had negative fatty acid content trends (mg EPA + DHA/g dry weight) in primary producers and primary consumers, but not in secondary nor tertiary fish consumers. The massive biomass increment of fish led to increasing areal fatty acid content (kg EPA + DHA/ha) towards increasingly warmer, more productive lakes, but there were no significant trends in other trophic levels. Increasing temperature and productivity are shifting subarctic lake communities towards systems characterized by increasing dominance of cyanobacteria and cyprinid fish, although decreasing quality in terms of EPA + DHA content was observed only in phytoplankton, zooplankton and profundal benthos.

A priori knowledge of fatty acid modifications in consumers is essential for studies using fatty acids as biomarkers. We investigated fatty acid metabolism and possible modification pathways in benthic invertebrate Chironomus riparius larvae (Diptera). We conducted diet manipulation experiments using natural food sources (two chlorophyte algae, a diatom and a non-toxic cyanobacterium). We also did a diet-switch experiment on two different resources, fish food flakes TetraMin ® and cyanobacterium Spirulina , to study fatty acid turnover in Chironomus . Results of the diet manipulation experiments indicate that Chironomus larvae have a strong tendency to biosynthesize 20:5n-3 and 20:4n-6 from precursor fatty acids, and that the dietary availability of polyunsaturated fatty acids (PUFA) does not control larval growth. Fatty acid modifications explain why low dietary availability of PUFA did not significantly limit growth. This has ecologically relevant implications on the role of benthic chironomids in conveying energy to upper trophic level consumers. A diet-switch experiment showed that the turnover rate of fatty acids in Chironomus is relatively fast––a few days. The compositional differences of algal diets were large enough to separate Chironomus larvae into distinct groups even if significant modification of PUFA was observed. In summary, fatty acids are excellent dietary biomarkers for Chironomus , if modifications of PUFA are considered, and will provide high-resolution data on resource use. This article is part of the theme issue ‘The next horizons for lipids as ‘trophic biomarkers': evidence and significance of consumer modification of dietary fatty acids'.

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 littoral macroinvertebrates, only Lymnaea (Gastropoda), Eurycercus (Cladocera), and Gammarus (Amphipoda) contained a low amount of DHA, whereas most littoral invertebrate taxa contained moderate amounts of eicosapentaenoic acid (EPA). The difference in DHA content may explain why so many generalist fish shift their diet to pelagic zooplankton at their peak abundance in mid/late‐summer. Meanwhile, the differences in EAA content between pelagic zooplankton and littoral invertebrates were much lower than for EFA suggesting a wider availability of EAA in subarctic lakes, except for methionine. In the studied subarctic lakes, EFA and EAA variation in consumers was more related to taxon‐specific than lake‐specific characteristics. This indicates that climate‐induced changes in the abundance and community structure of zooplankton vs. littoral macroinvertebrates will be important parameters in determining the availability of EFA and EAA to juvenile fish, and potentially fish production.

. Influence of littoral periphyton on whole-lake metabolism relates to littoral vegetation in humic lakes. Ecology, 98 (12), 3074-3085. doi:10.1002 Abstract. The role of littoral habitats in lake metabolism has been underrated, especially in humic lakes, based on an assumption of low benthic primary production (PP) due to low light penetration into water. This assumption has been challenged by recent recognition of littoral epiphyton dominance of whole-lake PP in a small highly humic lake and of epiphyton as an important basal food source for humic lake biota. However, as these studies have mostly concerned single lakes, there is a need to test their wider generality. We studied the whole-lake PP and community respiration (CR) in eight small humic lakes in southern Finland during July 2015 using 14 C incorporation to measure pelagic PP and the changes in dissolved inorganic carbon in light and dark in situ incubations to measure CR and littoral PP by epiphyton. Changes in O 2 concentration in both pelagic and littoral surface water were measured periodically from each lake and, additionally, continuously with a data logger from one lake during the study period. The results revealed that the littoral dominated whole-lake net primary production (NPP) in five of the eight lakes, which was supported by observed O 2 supersaturation in the littoral surface water in most of the lakes. Calculated pelagic:littoral ratios by area correlated negatively with both littoral NPP and littoral contribution to whole-lake NPP. Moreover, there was a significant positive relationship between littoral proportion of whole-lake NPP and the fraction of lake surface area covered by littoral aquatic vegetation. This demonstrates that increased aquatic littoral vegetation cover increases the overall importance of the littoral to whole-lake PP in highly humic lakes. Littoral NPP also correlated strongly with littoral O 2 saturation, and the continuously measured O 2 revealed substantial temporal variation in O 2 saturation, particularly in the littoral zone. Whole-lake gross primary production:community respiration (GPP:CR) ratios revealed that accounting for littoral metabolism produced a marked shift towards lake metabolic balance, although all the eight lakes remained net heterotrophic. This study emphasizes that littoral metabolism needs to be accounted for when estimating whole-lake C fluxes in all lakes, even in highly colored humic waters.