Ann-Kristin Bergström

Shifts in Lake N:P Stoichiometry and Nutrient Limitation Driven by Atmospheric Nitrogen Deposition

Nitrogen Overload The cycling of essential nutrients in terrestrial ecosystems has been altered by human activities. Elser et al. (p. 835) report a comparative analysis of lakes in Norway, Sweden, and in the United States that suggests that this is also true in aquatic ecosystems such as lakes. Deposition of anthropogenically derived atmospheric nitrogen controls whether N or P is growth-limiting for phytoplankton. Under elevated conditions of atmospheric N inputs, lake phytoplankton become consistently P-limited because the N:P ratio is strongly distorted. This is in contrast to conditions of low N deposition when lake phytoplankton are N-limited. These effects are even observed in remote lakes, demonstrating the indirect yet wide-ranging effects of humans on global food webs. Deposition of anthropogenically derived nitrogen can cause phosphorus to become the limiting nutrient of lake phytoplankton. Human activities have more than doubled the amount of nitrogen (N) circulating in the biosphere. One major pathway of this anthropogenic N input into ecosystems has been increased regional deposition from the atmosphere. Here we show that atmospheric N deposition increased the stoichiometric ratio of N and phosphorus (P) in lakes in Norway, Sweden, and Colorado, United States, and, as a result, patterns of ecological nutrient limitation were shifted. Under low N deposition, phytoplankton growth is generally N-limited; however, in high–N deposition lakes, phytoplankton growth is consistently P-limited. Continued anthropogenic amplification of the global N cycle will further alter ecological processes, such as biogeochemical cycling, trophic dynamics, and biological diversity, in the world’s lakes, even in lakes far from direct human disturbance.

ALLOCHTHONOUS ORGANIC CARBON AND PHYTOPLANKTON/BACTERIOPLANKTON PRODUCTION RELATIONSHIPS IN LAKES

Humic lakes with high inputs of allochthonous dissolved organic carbon have a pelagic food chain that, to a large extent, is based on bacterioplankton energy mobilization from allochthonous organic carbon compounds. This is in contrast to clear lakes in which total pelagic production is based mainly on phytoplankton photosynthesis. The energy economy in humic lakes may be less efficient than in clear lakes, because it is likely that one more link is included in the food chain. Lake data from Scandinavia and North America demonstrate that shifts between food chains based on heterotrophic production and food chains based on primary production can take place at moderate increases or decreases in the concentration of dissolved organic carbon from allochthonous sources. Large variations in the loading of allochthonous organic carbon (e.g., due to climatic variations) may have considerable effects on the biostructure and productivity of lakes.

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.

The use of TN:TP and DIN:TP ratios as indicators for phytoplankton nutrient limitation in oligotrophic lakes affected by N deposition

The stoichiometric composition of lake water chemistry affects nutrient limitation among phytoplankton. I show how TN:TP and DIN:TP ratios vary in oligotrophic lakes of Europe and the USA affected by different amounts of N deposition, and evaluate whether the DIN:TP ratio is a better indicator than the TN:TP ratio for discriminating between N and P limitation of phytoplankton. Data were compiled from boreal and low to high alpine lakes, and comprise epilimnetic lake water chemistry data (106 lakes) and results from short-term nutrient bioassay experiments (28 lakes). A large share (54%) of the oligotrophic lakes in the study had low TN:TP mass ratios (<25). DIN:TP ratios showed higher variability than TN:TP ratios. Variability in DIN:TP ratios was related to N deposition, but also to catchment characteristics. Data from short-term bioassay experiments with separate addition of N and P showed that the DIN:TP ratio was a better indicator than the TN:TP ratio for N and P limitation of phytoplankton. Phytoplankton shift from N to P limitation when DIN:TP mass ratios increase from 1.5 to 3.4. High DIN:TP ratios, indicating P limitation of phytoplankton, were generally found in alpine lakes with low to moderate N deposition and in boreal lakes with high to very high amounts of N deposition.

Bacterioplankton Production in Humic Lake Ortrasket in Relation to Input of Bacterial Cells and Input of Allochthonous Organic Carbon

A bstractIn order to compare riverine bacteria input with lake water bacterial production and grazing loss with output loss, a bacterial cell budget was constructed for humic Lake Örträsket in northern Sweden. The riverine input of bacterial cells in 1997 represented 29% of the number of bacterial cells produced within the layer of the lake affected by inlet water. A large share of the in situ lake bacterial production was consumed by grazers, mainly flagellates, which stresses the importance of bacteria as energy mobilizers for the pelagic food web in the lake. The bacterial production in Lake Örträsket, which is almost entirely dependent on humic material as an energy source, was clearly stimulated by high flow episodes which brought high amounts of little degraded material into the lake. During base flow condition the bacterial production in the inlet rivers was high, which led to an input of more degraded material to the lake. This material did not stimulate the lake bacterial production. Internal factors that determined the utilization of the allochthonous DOC in the lake were the retention time and the exposure to light and high temperatures. Thus, the potential for in situ production of bacteria in Lake Örträsket was to a large extent a function of how precipitation and runoff conditions affected terrestrial losses and river transport of humic material.

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.

Bacterial grazing by phagotrophic phytoflagellates in a deep humic lake in northern Sweden

Bacterial grazing was measured from June to August 1995 in Lake Ortrasket, a deep brown-water lake in northern Sweden. Mixotrophic chrysophytes were the dominating bacterivores at all times, grazin ...

Terrestrial organic matter input suppresses biomass production in lake ecosystems

Terrestrial ecosystems export large amounts of organic carbon (t-OC) but the net effect of this OC on the productivity of recipient aquatic ecosystems is largely unknown. In this study of boreal lakes, we show that the relative contribution of t-OC to individual top consumer (fish) biomass production, and to most of their potential prey organisms, increased with the concentration of dissolved organic carbon (DOC; dominated by t-OC sources) in water. However, the biomass and production of top consumers decreased with increasing concentration of DOC, despite their substantial use (up to 60%) of t-OC. Thus, the results suggest that although t-OC supports individual consumer growth in lakes to a large extent, t-OC input suppresses rather than subsidizes population biomass production.

Community composition of bacterioplankton and cell transport in lakes in two different drainage areas

Abstract.We investigated the influence of bacteria imported from the drainage area on the bacterioplankton communities in lakes. Six boreal humic lakes situated in two adjacent drainage areas were investigated. Budgets of cell transport were constructed by using a combination of hydrological and microbiological methods. Community composition of bacterioplankton in the lakes was determined by denaturing gradient gel electrophoresis (DGGE) of 16S rDNA. The results show that bacterial import can have been an important factor shaping the composition of bacterioplankton communities in the lakes, despite that the lakes were sampled at low flow occasions. Consequently, there were larger similarities in the composition of bacterioplankton between lakes within the same drainage area than between lakes in different drainage areas.

Intraspecific Autochthonous and Allochthonous Resource Use by Zooplankton in a Humic Lake during the Transitions between Winter, Summer and Fall

Seasonal patterns in assimilation of externally produced, allochthonous, organic matter into aquatic food webs are poorly understood, especially in brown-water lakes. We studied the allochthony (share biomass of terrestrial origin) in cladoceran, calanoid and cyclopoid micro-crustacean zooplankton from late winter to fall during two years in a small humic lake (Sweden). The use of allochthonous resources was important for sustaining a small population of calanoids in the water column during late winter. However, in summer the calanoids shifted to 100% herbivory, increasing their biomass several-fold by making efficient use of the pelagic primary production. In contrast, the cyclopoids and cladocerans remained at high levels of allochthony throughout the seasons, both groups showing the mean allochthony of 0.56 (range in mean 0.17-0.79 and 0.34-0.75, for the respective group, depending on model parameters). Our study shows that terrestrial organic matter can be an important resource for cyclopoids and cladocerans on an annual basis, forming a significant link between terrestrial organic matter and the higher trophic levels of the food web, but it can also be important for sustaining otherwise herbivorous calanoids during periods of low primary production in late winter.