Tom Andersen

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.

Ecological thresholds and regime shifts: approaches to identification

There is an apparent gap between the prominence of present theoretical frameworks involving ecological thresholds and regime shifts, and the paucity of efforts to conduct simple tests and quantitative inferences on the actual appearance of such phenomena in ecological data. A wide range of statistical methods and analytical techniques are now available that render these questions tractable, some of them even dating back half a century. Yet, their application has been sparse and confined within a narrow subset of cases of ecological regime shifts. Our objective is to raise awareness on the range of techniques available, and to their principles and limitations, to promote a more operational approach to the identification of ecological thresholds and regime shifts.

REDUCED LIGHT INCREASES HERBIVORE PRODUCTION DUE TO STOICHIOMETRIC EFFECTS OF LIGHT/NUTRIENT BALANCE

Ecological common sense says that decreased solar energy should reduce herbivore production because of reduced energy flow through primary producers. However, a field experiment in a phosphorus-limited lake showed that production of zooplankton herbivores was increased by shading. This paradoxical outcome was caused by a decoupling of producer carbon fixation and nutrient uptake under high light that reduced food quality for herbivores. At low nutrient supplies, shading increased nutrient contents relative to carbon within algal food, outweighing effects of decreased primary production. Thus, light/ nutrient balance affects the degree of mismatch between primary producers and herbivores in nature, which in turn influences mass-transfer efficiencies along food chains. To predict how energy transfer efficiency and biological interactions will respond to perturbations, it is essential to take into account changes in light/nutrient balance and its effects on the stoichiometry of autotroph-herbivore interactions.

Trends in Surface Water Acidification in Europe and North America (1989–1998)

During the last 20 years, emission reductions in Europe and North America have resulted in decreased atmospheric S-deposition of up to 50%, while N-deposition has stayed almost constant. Data from 98 ICP Waters sites were tested for trends in concentrations of major chemical components for the 10-year period 1989-1998 using the nonparametric seasonal Kendall test. The sites were grouped into regions and types for meta-analysis. All of the regions had highly significant downward trends in SO42−* concentrations. Nitrate concentrations, on the other hand, show no regional patterns of change. Concentrations of base cations declined in most regions. All regions showed tendencies of increasing DOC. The low ANC sites showed the largest rates of recovery. Neither the high NO3− or low NO3− groups of sites exhibited significant trends in NO3− concentrations. Alpine (non-forested) sites show clear and consistent signals of recovery in ANC and pH, and appropriate (relative to SO42−* trends) rates of base cation decline.

Factors influencing species richness in lacustrine zooplankton

Frequent dispersal events are expected to elevate local species richness in island-like habitats such as lakes. However, the importance of dispersal can be hard to evaluate if other factors cause large background variability in species composition and richness. In this paper, we review empirical studies on ecological factors known or expected to influence species richness in zooplankton communities of inland lakes. We then present summaries of two recent case studies. Our objectives are twofold: we first look for effects of biotic interactions on species richness and species composition, and then evaluate whether the expected effects of dispersal are likely to be detected on a background of large variability caused by other ecological factors and interactions. Species richness within lakes appears to be primarily controlled by factors related to lake size, lake productivity, water quality, and fish predation levels. One case study indicated a slight, but significant, positive effect of lake density and lake area in the surrounding landscape on species richness, suggesting that frequent dispersal events may enhance species richness. This local variation in species richness is superimposed on regional variation in species pools.

Pelagic food webs and eutrophication of coastal waters : Impact of grazers on algal communities

This review considers theoretical and empirical aspects of the role of grazers in the pelagic food web. We discuss how grazers may affect eutrophication development in coastal waters both through the direct effects of differential prey selectivity on the composition of the algal community, and through the indirect effects of nutrient sequestration and regeneration on the pelagic nutrient regime. We identify dominance of oligotrich ciliates or calanoid copepods on the primary herbivore level as two major food web configurations with different overall impacts on both algal communities and pelagic nutrient retention. Literature data on numerical and functional responses are compiled to make a quantitative assessment of competitive and predatory interactions between ciliates and copepods.

Regional species pools control community saturation in lake phytoplankton

Recent research has highlighted that positive biodiversity–ecosystem functioning relationships hold for all groups of organisms, including microbes. Yet, we still lack understanding regarding the drivers of microbial diversity, in particular, whether diversity of microbial communities is a matter of local factors, or whether metacommunities are of similar importance to what is known from higher organisms. Here, we explore the driving forces behind spatial variability in lake phytoplankton diversity in Fennoscandia. While phytoplankton biovolume is best predicted by local phosphorus concentrations, phytoplankton diversity (measured as genus richness, G) only showed weak correlations with local concentrations of total phosphorus. By estimating spatial averages of total phosphorus concentrations on various scales from an independent, spatially representative lake survey, we found that close to 70 per cent of the variability in local phytoplankton diversity can be explained by regionally averaged phosphorus concentrations on a scale between 100 and 400 km. Thus, the data strongly indicate the existence of metacommunities on this scale. Furthermore, we show a strong dependency between lake productivity and spatial community turnover. Thus, regional productivity affects beta-diversity by controlling spatial community turnover, resulting in scale-dependent productivity-diversity relationships. As an illustration of the interaction between local and regional processes in shaping microbial diversity, our results offer both empirical support and a plausible mechanism for the existence of common scaling rules in both the macrobial and the microbial worlds. We argue that awareness of regional species pools in phytoplankton and other unicellular organisms may critically improve our understanding of ecosystems and their susceptibility to anthropogenic stressors.

Heavy Metal Surveys in Nordic Lakes; Concentrations, Geographic Patterns and Relation to Critical Limits

Abstract In the autumn of 1995, coordinated national lake surveys were conducted in the Nordic countries, including Russian Kola. The 11 metals (Pb, Cd, As, Zn, Cu, Ni, Co, Fe, Mn, Cr, V) investigated in nearly 3000 lakes have generally low concentrations and distinct geographical patterns. Direct and indirect influence of long-range transported air pollution is the major important factor for distribution of Pb, Cd, Zn and to a certain degree Co. Total organic carbon (TOC) concentrations in lakes are important for Fe and Mn but also to a certain degree for As, Cr and V. Bedrock geology is the major controlling factor for Cu and Ni, with the exception of areas around the smelters in the Kola peninsula, where the Cu and Ni concentrations in lakes are very high due to local airborne pollution. Bedrock and surficial geology is also an important factor for controlling the concentrations of As, Co, Cr and V. The results indicate that heavy metal pollution in lakes is a minor ecological problem on a regional scale in the Nordic countries.

DYNAMIC ENERGY BUDGET REPRESENTATIONS OF STOICHIOMETRIC CONSTRAINTS ON POPULATION DYNAMICS

Metabolism, and thus population dynamics, can be limited by energy, carbon, nitrogen, and/or other nutrients. This is due to homeostasis, the relatively constant composition of biomass. Yet growth-rate-dependent changes in the composition of biomass do exist. The dynamic energy budget (DEB) theory provides the framework to deal with these simultaneous limitations and stoichiometric restrictions. We illustrate the application with three examples. First, we discuss simple single-species growth of a chemolithoautotroph to illustrate the interactions between nutrients and substrates in growth. We show how the macrochemical reaction equation with variable yield coefficients can be decomposed in a number of subprocesses with constant yield coefficients. We then discuss a simple predator– prey system, where nutrients are accumulated in the prey, which no longer have a constant composition of biomass. The implication is a varying conversion efficiency from prey to predator, with consequences for qualitative aspec...

Increased risk of phosphorus limitation at higher temperatures for Daphnia magna.

Invertebrate herbivores frequently face growth rate constraints due to their high demands for phosphorus (P) and nitrogen (N). Temperature is a key modulator of growth rate, yet the interaction between temperature and P limitation on somatic growth rate is scarcely known. To investigate this interaction, we conducted a study on the somatic growth rate (SGR) of the cladoceran Daphnia magna, known to be susceptible to P-limitation. We determined the SGR across a broad range of dietary P content of algae (carbon (C):P ratios (125–790), and at different temperatures (10–25°C). There was a strong impact of both temperature and C:P ratio on the SGR of D. magna, and also a significant interaction between both factors was revealed. The negative effect of dietary C:P on growth rate was reduced with decreased temperature. We found no evidence of P limitation at lowest temperature, suggesting that enzyme kinetics or other measures of food quality overrides the demands for P to RNA and protein synthesis at low temperatures. These findings also indicate an increased risk of P limitation and thus reduced growth efficiency at high temperatures.