Lakes play a significant role in the global carbon cycle where inputs from watersheds and primary production are either stored in sediments or lost to the atmosphere through respiration. Climate change is anticipated to increase atmospheric losses as water overlying sediments warms, thus reducing carbon storage. Lakes worldwide, however, are not only warming but are also losing transparency through eutrophication or browning. The synergistic result is that heat is trapped in the surface layers of more colored lakes, which in turn isolates colder bottom waters and sediments experience longer periods without oxygen. This bottom-water cooling increases overall carbon storage by reducing aerobic respiration, but stimulates methane production due to prolonged anoxia, thus potentially increasing the overall global warming potential of lakes. AbstractIn this article, we challenge the notion that global warming stimulates organic matter mineralization and increases greenhouse gas emissions in lakes via direct temperature effects. We show that the interactive effects of warming and transparency loss due to eutrophication or browning overrides atmospheric warming alone. Thermal shielding enables a longer and more stable stratification that results in bottom-water cooling, prolonged anoxia, and enhanced carbon preservation in a large proportion of global lakes. These effects are strongest in shallow lakes where an additional burial of 4.5 Tg C yr −1 increases current global estimates by 9%. Despite more burial, the net global warming potential of lakes will increase via enhanced methane production, related to prolonged periods of anoxia, rather than warming. Our understanding of how whole-lake carbon cycling responds to climate change needs revision, as the synergistic influence of warming and transparency loss has much broader ecosystem level functional consequences.

Mixing regime and CO 2 availability may control cyanobacterial blooms in polymictic lakes, but the underlying mechanisms still remain unclear. We integrated detailed results from a natural experiment comprising an average-wet year (2011) and one with heat waves (2012), a long-term meteorological dataset , historical phosphorus concentrations and sedimentary pigment records, to determine the mechanistic controls of cyanobacterial blooms in a eutrophic polymictic lake. Intense warming in 2012 was associated with: 1) increased stability of the water column with buoyancy frequencies exceeding 40 cph at the surface, 2) high phytoplankton biomass in spring (up to 125 mg WW L -1 ), 3) reduced downward transport of heat and 4) depleted epilimnetic CO 2 concentrations. CO 2 depletion was maintained by intense uptake by phytoplankton (influx up to 30 mmol m -2 d -1 ) in combination with reduced, internal and external, carbon inputs during dry, stratified periods. These synergistic effects triggered bloom of buoyant cyanobacteria (up to 300 mg WW L -1 ) in the hot year. Complementary evidence from polynomial regression modelling using historical data and pigment record revealed that warming explains 78% of the observed trends in cyanobacterial biomass, whereas historical phosphorus concentration only 10% thereof. Together the results from the natural experiment and the longterm record indicate that effects of hotter and drier climate are likely to increase water column stratification and decrease CO 2 availability in eutrophic polymictic lakes. This combination will catalyze blooms of buoyant cyanobacteria.

Abstract. Permafrost thaw lakes (thermokarst lakes) are widely distributed across the northern landscape, and are known to be biogeochemically active sites that emit large amounts of carbon to the atmosphere as CH4 and CO2. However, the abundance and composition of the photosynthetic communities that fix CO2 have been little explored in this ecosystem type. In order to identify the major groups of phototrophic organisms and their controlling variables, we sampled 12 permafrost thaw lakes along a permafrost degradation gradient in northern Québec, Canada. Additional samples were taken from five rock-basin reference lakes in the region to determine if the thaw lakes differed in limnological properties and phototrophs. Phytoplankton community structure was determined by high-performance liquid chromatography analysis of their photoprotective and photosynthetic pigments, and autotrophic picoplankton concentrations were assessed by flow cytometry. One of the black-colored lakes located in a landscape of rapidly degrading palsas (permafrost mounds) was selected for high-throughput 18S rRNA sequencing to complement conclusions based on the pigment and cytometry analyses. The results showed that the limnological properties of the thaw lakes differed significantly from the reference lakes, and were more highly stratified. However, both waterbody types contained similarly diverse phytoplankton groups, with dominance of the pigment assemblages by fucoxanthin-containing taxa, as well as chlorophytes, cryptophytes and cyanobacteria. Chlorophyll a concentrations (Chl a) were correlated with total phosphorus (TP), and both were significantly higher in the thaw lakes (overall means of 3.3 µg Chl a L−1 and 34 µg TP L−1) relative to the reference lakes (2.0 µg Chl a L−1 and 8.2 µg TP L−1). Stepwise multiple regression of Chl a against the other algal pigments showed that it was largely a function of alloxanthin, fucoxanthin and Chl b (R2 =  0.85). The bottom waters of two of the thaw lakes also contained high concentrations of bacteriochlorophyll d, showing the presence of green photosynthetic sulphur bacteria. The molecular analyses indicated a relatively minor contribution of diatoms, while chrysophytes, dinoflagellates and chlorophytes were well represented; the heterotrophic eukaryote fraction was dominated by numerous ciliate taxa, and also included Heliozoa, Rhizaria, chytrids and flagellates. Autotrophic picoplankton occurred in biovolume concentrations up to 3.1  ×  105 µm3 picocyanobacteria  mL−1 and 1.9  × 106 µm3 picoeukaryotes mL−1, with large variations among lakes. Both groups of picophytoplankton were positively correlated with total phytoplankton abundance, as measured by Chl a; picocyanobacteria were inversely correlated with dissolved organic carbon, while picoeukaryotes were inversely correlated with conductivity. Despite their net heterotrophic character, subarctic thaw lakes are rich habitats for diverse phototrophic communities.

Harmful cyanobacterial blooms are an increasing problem at many locations throughout the world but are rarely reported in aquatic habitats at high latitudes. Shallow lakes are a major feature of northern permafrost landscapes and are likely to experience large‐scale changes in their limnological properties in the future as a consequence of climate warming. In the present study, we addressed the question of what preconditions would be necessary to stimulate the growth and dominance of bloom‐forming cyanobacteria in northern fresh waters. We analysed the summer phytoplankton of 18 lakes on eroding permafrost (thaw lakes) and on glacier‐scoured rock (rock basin lakes) in subarctic Quebec, Canada, to determine their phytoplankton community structure and the biomass contribution of cyanobacteria. This survey was complemented with an incubation experiment to evaluate the direct warming and indirect phosphorus (P) enrichment effects of climate change on cyanobacterial bloom development. All lakes contained diverse phytoplankton communities, often dominated by chrysophytes, dinoflagellates and chlorophytes. Cyanobacteria were present in all waterbodies, but their contribution to the total community biovolume was highly variable (mean of 8.7%, range 0.1%–47%). Cyanobacterial community biovolumes correlated positively with surface water temperatures, and negatively with dissolved organic carbon, soluble reactive phosphorus, iron and manganese concentrations in the surface waters. Phosphorus enrichment of water from a thaw lake resulted in a fourfold increase of chlorophyll a (Chl‐a) and an increase in the cyanobacterial pigments echinenone and zeaxanthin. The phytoplankton counts showed that there was a sharp decrease in diversity (expressed as decline of the Shannon–Wiener index from 1.69 to 0.16), accompanied by a shift to cyanobacterial dominance, notably by the heterocystous, potentially toxic species Dolichospermum cf. planctonicum. Increased temperature led to an initial doubling of cyanobacterial biovolume, followed by the development of a chrysophyte bloom. Combined warming and P enrichment led to reduced phytoplankton biodiversity, with a community composed of cyanobacteria and chrysophytes. There was also a pronounced response by the picophytoplankton community; picocyanobacteria were strongly stimulated by P enrichment, while picoeukaryotes increased in response to warming. The current inoculum levels of cyanobacteria in subarctic lakes and their responsiveness to temperature and phosphorus indicate the potential for an abrupt increase in their abundance, accompanied by a decrease in phytoplankton diversity. Ongoing climate change will increase the risk of noxious cyanobacterial blooms in northern lakes and ponds, with potentially negative consequences for higher trophic levels.

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Climate change is proceeding rapidly at high northern latitudes and may have a variety of direct and indirect effects on aquatic food webs. One predicted effect is the potential shift in phytoplankton community structure towards increased cyanobacterial abundance. Given that cyanobacteria are known to be a nutritionally poor food source, we hypothesized that such a shift would reduce the efficiency of feeding and growth of northern zooplankton. To test this hypothesis, we first isolated a clone of Daphnia pulex from a permafrost thaw pond in subarctic Québec, and confirmed that it was triploid but otherwise genetically similar to a diploid, reference clone of the same species isolated from a freshwater pond in southern Québec. We used a controlled flow-through system to investigate the direct effect of temperature and indirect effect of subarctic picocyanobacteria (Synechococcus) on threshold food concentrations and growth rate of the high latitude clone. We also compared the direct effect of temperature on both Daphnia clones feeding on eukaryotic picoplankton (Nannochloropsis). The high latitude clone had a significantly lower food threshold for growth than the temperate clone at both 18 and 26°C, implying adaptation to lower food availability even under warmer conditions. Polyunsaturated fatty acids were present in the picoeukaryote but not the cyanobacterium, confirming the large difference in food quality. The food threshold for growth of the high latitude Daphnia was 3.7 (18°C) to 4.2 (26°C) times higher when fed Synechococcus versus Nannochloropsis, and there was also a significant negative effect of increased temperature and cyanobacterial food on zooplankton fatty acid content and composition. The combined effect of temperature and food quality on the performance of the high latitude Daphnia was greater than their effects added separately, further indicating the potentially strong indirect effects of climate warming on aquatic food web processes.

Abstract. Permafrost thaw lakes (thermokarst lakes) are widely distributed across the northern landscape, and are known to be biogeochemically active sites that emit large amounts of carbon to the atmosphere as CH4 and CO2. However, the abundance and composition of the photosynthetic communities that consume CO2 have been little explored in this ecosystem type. In order to identify the major groups of phototrophic organisms and their controlling variables, we sampled 12 permafrost thaw lakes along a permafrost degradation gradient in northern Québec, Canada. Additional samples were taken from 5 rock-basin reference lakes in the region to determine if the thaw waters differed in limnological properties and phototrophs. Phytoplankton community structure was determined by high performance liquid chromatography analysis of their photoprotective and photosynthetic pigments, and autotrophic picoplankton concentrations were assessed by flow cytometry. One of the black colored lakes located in a andscape of rapidly degrading palsas (permafrost mounds) was selected for high-throughput 18S rRNA sequencing to help interpret the pigment and cytometry data. The results showed that the limnological properties of the thaw lakes differed significantly from the reference lakes, and were more highly stratified. However, both waterbody types contained similarly diverse phytoplankton groups, with dominance of the pigment assemblages by fucoxanthin-containing taxa, as well as chlorophytes, cryptophytes and cyanobacteria. Chlorophyll a concentrations (Chl a) were correlated with total phosphorus (TP), and both were significantly higher in the thaw lakes (overall means of 3.3 μg Chl a L−1 and 34 μg TP L−1) relative to the reference lakes (2.0 μg Chl a L−1 and 8.2 μg TP L−1). Stepwise multiple regression of Chl a against the other algal pigments showed that it was largely a function of lutein, fucoxanthin and peridinin (R2 = 0.78). The bottom waters of two of the thaw lakes also contained high concentrations of bacteriochlorophyll d, showing the presence of green photosynthetic sulphur bacteria. The molecular analyses indicated a relatively minor contribution of diatoms, while chrysophytes, dinoflagellates and chlorophytes were well represented; the heterotrophic eukaryote fraction was dominated by numerous ciliate taxa, and also included Heliozoa, Rhizaria, chytrids and flagellates. Autotrophic picoplankton occurred in cell concentrations up to 8.8 × 105 mL−1 (picocyanobacteria) and 4.6 × 105 mL−1 (picoeukaryotes). Both groups of picophytoplankton were positively correlated with total phytoplankton abundance, as measured by Chl a; picocyanobacteria were inversely correlated with dissolved organic carbon, while picoeukaryotes were correlated with conductivity. Despite their net heterotrophic character, subarctic thaw lakes are rich habitats for diverse phototrophic communities.