Dag O. Hessen

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.

CARBON SEQUESTRATION IN ECOSYSTEMS: THE ROLE OF STOICHIOMETRY

The fate of carbon (C) in organisms, food webs, and ecosystems is to a major extent regulated by mass-balance principles and the availability of other key nutrient elements. In relative terms, nutrient limitation implies excess C, yet the fate of this C may be quite different in autotrophs and heterotrophs. For autotrophs nutrient limitation means less fixation of inorganic C or excretion of organic C, while for heterotrophs nutrient limitation means that more of ingested C will “go to waste” in the form of egestion or respiration. There is in general a mismatch between autotrophs and decomposers that have flexible but generally high C:element ratios, and consumers that have lower C:element ratios and tighter stoichiometric regulation. Thus, C-use efficiency in food webs may be governed by the element ratios in autotroph biomass and tend to increase when C:element ratios in food approach those of consumers. This tendency has a strong bearing on the sequestration of C in ecosystems, since more C will be di...

The impact of nutrient state and lake depth on top-down control in the pelagic zone of lakes: A study of 466 lakes from the temperate zone to the Arctic

Using empirical data from 466 temperate to arctic lakes covering a total phosphorus (TP) gradient of 2–1036 gL –1 , we describe how the relative contributions of resource supply, and predator control change along a nutrient gradient. We argue that (a) predator control on large-bodied zooplankton is unimodally related to TP and is highest in the most nutrient-rich and nutrient-poor lakes and generally higher in shallow than deep lakes, (b) the cascading effect of changes in predator control on phytoplankton decreases with increasing TP, and (c) these general patterns occur with significant variations—that is, the predation pressure can be low or high at all nutrient levels. A quantile regression revealed that the median share of the predatorsensitive Daphnia to the total cladoceran biomass was significantly related unimodally to TP, while the 10% and 90% percentiles approached 0 and 100%, respectively, at all TP levels. Moreover, deep lakes (more than 6 m) had a higher percentage of

Nutrient Element Limitation of Zooplankton Production

Freshwater crustacean zooplankton species show pronounced interspecific variability but approach intraspecific homeostasis in their stoichiometry. It is especially interesting that members of the dominating herbivores in freshwater systems, such as Daphnia species, show consistently high P:C ratios. Sestonic P:C ratios in epilimnetic waters are usually far below that encountered in the zooplankton, which suggests that grazers may approach a direct phosphorus limitation in many localities. This view was supported by a regional study of 47 lakes, in which most independent variables including phytoplankton biomass and chlorophyll were scarcely correlated with zooplankton biomass but particulate P (>0.45 μm) could explain a major part of the variance of zooplankton biomass. Of the various taxa, daphnids gave the best correlation with particulate P, cyclopids and rotifers the worst. These observations suggest that phytoplankton biomass per se may not be the major determinant of zooplankton biomass but that the nutritional status of phytoplankton (P cell quotas) rather than biomass in terms of carbon is most important, or that zooplankton suffer a direct P limitation in many lakes. The latter view is supported by model predictions given in this article. Also, both freshwater and marine copepod grazers could correspondingly be growth limited by an N:C ratio of ingested food that is too low The possibility of direct nutrient element limitation has wide implications for the regulation of herbivore production, competition, and overall cycling of nutrients in epilimnetic waters.

LIGHT, NUTRIENTS, AND P:C RATIOS IN ALGAE: GRAZER PERFORMANCE RELATED TO FOOD QUALITY AND QUANTITY

Continuous cultures of the green algae Selenastrum capricornutum were response to self-shading. In that case, our results indicate that light adaptation not only involves a cost in terms of increased chlorophyll synthesis, but also in terms of increased P demands. This provides new insight not only into the physiological regulation of C and P uptake in algae, but it could also explain deviations from the Redfield ratio. The growth of juvenile Daphnia magna fed S. capricornutum from the different light and phosphorus treatments was studied in a series of short-term assays (7 d) covering a gradient of food concentrations. The response of Daphnia growth rate. along this quantity (0.5-5.0 mg C/L) and quality (0.5-12 Rg atomic P.(mg atomic C)-1) gradient gave a close fit to a double hyperbola model. Changes in elemental ratios of the algae were reflected in the growth rate of Daphnia, such that up to 40% reduction in its growth rate could be attributed to increased C:P ratios. This study demonstrates that the physiological responses of phototrophs in terms of chlorophyll content and elemental composition depend strongly on ambient light and nutrient regimes. It also confirms that these patterns can yield con- trasting responses on herbivore growth responses along the food quantity and quality axes.

Metabolic stoichiometry and the fate of excess carbon and nutrients in consumers

Animals encountering nutritionally imbalanced foods should release elements in excess of requirements in order to maintain overall homeostasis. Quantifying these excesses and predicting their fate is, however, problematic. A new model of the stoichiometry of consumers is formulated that incorporates the separate terms in the metabolic budget, namely, assimilation of ingested substrates and associated costs, protein turnover, other basal costs, such as osmoregulation, and the use of remaining substrates for production. The model indicates that release of excess C and nonlimiting nutrients may often be a significant fraction of the total metabolic budget of animals consuming the nutrient‐deficient forages that are common in terrestrial and aquatic systems. The cost of maintenance, in terms of not just C but also N and P, is considerable, such that food quality is important even when intake is low. Many generalist consumers experience short‐term and unpredictable fluctuations in their diets. Comparison of model output with data for one such consumer, Daphnia, indicates that mechanisms operating postabsorption in the gut are likely the primary means of regulating excess C, N, and P in these organisms, notably respiration decoupled from biochemical or mechanical work and excretion of carbon and nutrients. This stoichiometrically regulated release may often be in organic rather than inorganic form, with important consequences for the balance of autotrophic and heterotrophic processes in ecosystems.

Filtering structures and particle size selection in coexisting Cladocera

SummaryFive commonly co-occurring cladocerans, Holopedium gibberum, Daphnia longispina, Ceriodaphnia quadrangula, Bosmina longispina, Diaphanosoma brachyurum and the calanoid copepod Eudiaptomus gracilis were fed monodisperse fluorescent latex beads (0.5, 1,5 μm) to reveal clearance rates and particle size selection. The results were compared to those obtained with labelled bacteria and yeast. All cladoceran species ingested the beads, and particle size selection varied from a strong predominance of the largest particles in Holopedium, to an almost non selective particle retention in Diaphanosoma. The results indicated a strong correlation between morphology of the filtering combs, measured by Scanning electron microscopy, and ability to retain the smallest particles.

Competitive trade-off strategies in ArcticDaphnia linked to melanism and UV-B stress

Different high-Arctic, freshwater localities at Ny-Ålesund, Svalbard (79dgN) were examined for their UV-absorbing properties, depth and presence of melanic Or non-melanic morphs of the planktonic crustaceanDaphnia pulex. Light regimes in two localities with each of these morphs were measured by using underwater spectroradiometer. Most localities have low absorbance of short-waved light, but no clearcut relationship between UV transparency and occurrence of melanic morphs was detected. Yet, in the laboratory, the melanic morph showed far lower growth rates, thus being competitively inferior to the non-melanic. Conversely, the melanic morph was more resistant to UV light, suggesting a trade-off between the metabolic tax paid for the melanin synthesis and its UV-protecting abilities, or a staggered growth capacity possibly owing to polyploidy. Frequency of melanic or non-melanic clones could thus be directly linked to ambient UV-B stress and serve as an indicator thereof, but the apparently extensive need for UV protection under the Arctic lighl regimes is still puzzling, and the role of melanism and polyploidy in these organisms cannot be considered finally settled.

Extrinsic and intrinsic controls of zooplankton diversity in lakes

Pelagic crustacean zooplankton were collected from 336 Norwegian lakes covering a wide range of latitude, altitude, lake area, mean depth, production (as chlorophyll a), and fish community structure. Mean zooplankton species richness during the ice-free season was generally low at high latitudes and altitudes. Further, lower species richness was recorded in western lakes, possibly reflecting constraints on migration and dispersal. However, despite obvious spatial limitations, geographic boundaries were only weak predictors of mean zooplankton richness. Similarly, lake surface area did not contribute positively to mean richness such as seen in other ecosystem surveys. Rather, intrinsic factors such as primary production and fish community (planktivore) structure were identified by regression analysis as the major predictors of zooplankton diversity, while a positive correlation was observed between species richness and total zooplankton biomass. However, in spite of a large number of variables included in this study, the predictive power of multiple regression models was modest (<50% variance explained), pointing to a major role for within-lake properties, as yet unidentified intrinsic forces, stochasticity, or dispersal as constraints on zooplankton diversity in these lakes.

Reduced Fitness of Daphnia magna Fed a Bt-Transgenic Maize Variety

Genetically modified (GM) maize expressing the Bt-toxin Cry1Ab (Bt-maize) was tested for effects on survival, growth, and reproduction of the water flea Daphnia magna, a crustacean arthropod commonly used as a model organism in ecotoxicological studies. In three repeated experiments, D. magna were fed 100% ground maize in suspension, using either GM or isogenic unmodified (UM) maize. D. magna fed GM-maize showed a significantly reduced fitness performance: The mortality was higher, a lower proportion of females reached sexual maturation, and the overall egg production was lower compared to D. magna fed UM isogenic maize. We conclude that the tested variety of Bt-maize and its UM counterpart do not have the same quality as food sources for this widely used model organism. The combination of a reduced fitness performance combined with earlier onset of reproduction of D. magna fed Bt-maize indicates a toxic effect rather than a lower nutritional value of the GM-maize.