Pelagic food webs of humic lakes show low short-term response to forest harvesting.

Forest harvest in the boreal zone can increase the input of terrestrial materials such as dissolved organic carbon (DOC) and nitrate (NO3- ) into nearby aquatic ecosystems, with potential effects on phytoplankton growth through enhanced nutrient (i.e., positive) or reduced light availability (i.e., negative), which may affect ecosystem productivity and consumer resource use. Here, we conducted forest clear-cutting experiments in the catchments of four small, humic, and nitrogen-limited unproductive boreal lakes (two controls and two clear-cut, 18% and 44% of area cut) with one reference and two impact years. Our aim was to assess the effects of forest clear-cutting on pelagic biomass production and consumer resource use. We found that pelagic biomass production did not change after two years of forest clear-cutting: Pelagic primary and bacterial production (PP, BP), PP:BP ratio, chl a, and seston carbon (seston C) were unaffected by clear-cutting; neither did tree harvest affect seston stoichiometry (i.e., N:phosphorus [P], C:P) nor induce changes in zooplankton resource use, biomass, or community composition. In conclusion, our findings suggest that pelagic food webs of humic lakes (DOC > 15 mg/L) might be resilient to a moderate form of forest clear-cutting, at least two years after tree removal, before mechanical site preparation (e.g., mounding, plowing) and when leaving buffer strips along lakes and incoming streams. Thus, pelagic food web responses to forest clear-cutting might not be universal, but could depend on factors such as the time scale, share of catchment logged, and the forest practices involved, including the application of buffer strips and site preparation.

Nitrogen effects on the pelagic food web are modified by dissolved organic carbon.

Global environmental change has altered the nitrogen (N) cycle and enhanced terrestrial dissolved organic carbon (DOC) loadings to northern boreal lakes. However, it is still unclear how enhanced N availability affects pelagic food web efficiency (FWE) and crustacean zooplankton growth in N limited boreal lakes. Here, we performed in situ mesocosm experiments in six unproductive boreal Swedish lakes, paired across a DOC gradient, with one lake in each pair fertilized with N (2011: reference year; 2012, 2013: impact years). We assessed how zooplankton growth and FWE were affected by changes in pelagic energy mobilization (PEM), food chain length (phytoplankton versus bacterial production based food chain, i.e. PP:BP), and food quality (seston stoichiometry) in response to N fertilization. Although PP, PEM and PP:BP increased in low and medium DOC lakes after N fertilization, consumer growth and FWE were reduced, especially at low DOC-potentially due to reduced phytoplankton food quality [increased C: phosphorus (P); N:P]. At high DOC, N fertilization caused modest increases in PP and PEM, with marginal changes in PP:BP and phytoplankton food quality, which, combined, led to a slight increase in zooplankton growth and FWE. Consequently, at low DOC (<12 mg L-1), increased N availability lowers FWE due to mismatches in food quality demand and supply, whereas at high DOC this mismatch does not occur, and zooplankton production and FWE may increase. We conclude that the lake DOC level is critical for predicting the effects of enhanced inorganic N availability on pelagic productivity in boreal lakes.

Phytoplankton response to whole lake inorganic N fertilization along a gradient in dissolved organic carbon.

Global change has increased inorganic nitrogen (N) and dissolved organic carbon (DOC; i.e., "browning") inputs to northern hemisphere boreal lakes. However, we do not know how phytoplankton in nutrient poor lake ecosystems of different DOC concentration respond to increased N availability. Here, we monitored changes in phytoplankton production, biomass and community composition in response to whole lake inorganic N fertilization in six boreal unproductive Swedish lakes divided into three lake pairs (control, N enriched) at three DOC levels (low, medium, high), with one reference year (2011) and 2 impact yr (2012, 2013). We found that phytoplankton biomass and production decreased with DOC concentration before N fertilization. Further, phytoplankton community composition also differed with respect to DOC, with a dominance of non-flagellated autotrophs at low DOC towards an increasing dominance of flagellated autotrophs with increased lake DOC concentration. The N fertilization increased phytoplankton biomass and production in all lakes, but did not affect phytoplankton community composition. However, the net response in biomass and production to N fertilization declined with increasing DOC, implying that the lake DOC concentration is critical in order to infer phytoplankton responses to N fertilization, and that the system switches from being primarily nutrient limited to becoming increasingly light limited with increased DOC concentration. In conclusion, our results show that browning will reduce phytoplankton production and biomass and influence phytoplankton community composition, whereas increased inorganic N loadings from deposition, forestry or other land use will primarily enhance phytoplankton biomass and production. Together, any change in the landscape that enhances inorganic N availability will increase phytoplankton production and biomass, but the positive effects of N will be much weaker or even neutralized in browner lakes as caused by light limitation.

Food web efficiency differs between humic and clear water lake communities in response to nutrients and light.

This study demonstrates that clear and humic freshwater pelagic communities respond differently to the same environmental stressors, i.e. nutrient and light availability. Thus, effects on humic communities cannot be generalized from existing knowledge about these environmental stressors on clear water communities. Small humic lakes are the most numerous type of lake in the boreal zone, but little is known about how these lakes will respond to increased inflows of nutrients and terrestrial dissolved organic C (t-DOC) due to climate change and increased human impacts. Therefore, we compared the effects of nutrient addition and light availability on pelagic humic and clear water lake communities in a mesocosm experiment. When nutrients were added, phytoplankton production (PPr) increased in both communities, but pelagic energy mobilization (PEM) and bacterial production (BP) only increased in the humic community. At low light conditions, the addition of nutrients led to increased PPr only in the humic community, suggesting that, in contrast to the clear water community, humic phytoplankton were already adapted to lower ambient light levels. Low light significantly reduced PPr and PEM in the clear water community, but without reducing total zooplankton production, which resulted in a doubling of food web efficiency (FWE = total zooplankton production/PEM). However, total zooplankton production was not correlated with PEM, PPr, BP, PPr:BP or C:nutrient stoichiometry for either community type. Therefore, other factors such as food chain length, food quality, ultra-violet radiation or duration of the experiment, must have determined total zooplankton production and ultimately FWE.

Lake secondary production fueled by rapid transfer of low molecular weight organic carbon from terrestrial sources to aquatic consumers.

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.

Effects of Additions of DOC on Pelagic Biota in a Clearwater System: Results from a Whole Lake Experiment in Northern Sweden.

An oligotrophic clearwater lake, initially characterized by a pronounced dominance of autotrophic phytoplankton and mostly by one species, the green alga Botryococcus, was subject to additions of dissolved organic carbon in the form of white sugar (sucrose) during two consecutive years. The hypothesis tested was that it is organic carbon per se, and not other possible effects of humic substances, that determines the differences in structure of the planktonic ecosystem between humic and clearwater lakes. The additions of DOC resulted in a significant increase in bacterial biomass and a decrease in the biomass of autotrophic phytoplankton. The biomass of mixotrophic and heterotrophic flagellates instead increased significantly, whereas no effects were found to propagate to higher trophic levels. As a result of the changes among biota, total planktonic biomass also decreased to a level typical of nearby humic lakes. We suggest that it is the carbon component of humic material and its utilization by bacterioplankton that determines the structure and function of the pelagic food web in humic lakes.