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.

Atmospheric nitrogen (N) deposition to lakes and watersheds has been increasing steadily due to various anthropogenic activities. Because such anthropogenic N is widely distributed, even lakes relatively removed from direct human disturbance are potentially impacted. However, the effects of increased atmospheric N deposition on lakes are not well documented. We examined phytoplankton biomass, the absolute and relative abundance of limiting nutrients (N and phosphorus [P]), and phytoplankton nutrient limitation in alpine lakes of the Rocky Mountains of Colorado (USA) receiving elevated (> 6 kg N x ha(-1) x yr(-1)) or low (< 2 kg N x ha(-1) x yr(-1)) levels of atmospheric N deposition. High-deposition lakes had higher NO3-N and total N concentrations and higher total N : total P ratios. Concentrations of chlorophyll and seston carbon (C) were 2-2.5 times higher in high-deposition relative to low-deposition lakes, while high-deposition lakes also had higher seston C:N and C:P (but not N:P) ratios. Short-term enrichment bioassays indicated a qualitative shift in the nature of phytoplankton nutrient limitation due to N deposition, as high-deposition lakes had an increased frequency of primary P limitation and a decreased frequency and magnitude of response to N and to combined N and P enrichment. Thus elevated atmospheric N deposition appears to have shifted nutrient supply from a relatively balanced but predominantly N-deficient regime to a more consistently P-limited regime in Colorado alpine lakes. This adds to accumulating evidence that sustained N deposition may have important effects on lake phytoplankton communities and plankton-based food webs by shifting the quantitative and qualitative nature of nutrient limitation.

Recently, data have emerged indicating that not only high food carbon:phosphorus (C:P) ratio but also low food C:P (P-rich food) can have negative effects on the growth of consumers. The shape of this "stoichiometric knife edge," however, is not yet well-documented, and the mechanisms underpinning it are not understood. Here we report the results of experiments using 3 species of Daphnia (D. magna, D. pulicaria, D. pulex) consuming the green alga Scenedesmus acutus with widely varying C:P ratios (from <50 to >1500 by atoms). The experiments were designed to (1) characterize the potential stoichiometric knife edge for each species, and (2) evaluate potential changes in feeding and respiration rates that may underpin the unimodal response to food C:P. All 3 Daphnia species grew more slowly when food C:P (atomic) exceeded ~250-300 but also when C:P was <120. Both high and low C:P foods were associated with increased respiration rates, indicating that the negative effects of food C:P imbalance at least partially involve increased metabolic costs of dealing with stoichiometrically imbalanced food. Feeding rate experiments indicated that, in contrast with limited previous data, animals generally increased their feeding rate on P-rich food. Overall, the "lower threshold elemental ratio" we identify here (~120) is surprisingly high, in an ecologically meaningful range, suggesting that negative effects of excessive food P content may play an under-recognized role in affecting Daphnia performance in P-rich lakes with low seston C:P ratio. Such effects also need to be incorporated into stoichiometrically explicit models of planktonic trophic interactions.

Cuatro Ciénegas Basin (CCB) is a desert ecosystem that hosts a large diversity of water bodies. Many surface waters in this basin have imbalanced nitrogen (N) to phosphorus (P) stoichiometry (total N:P > 100 by atoms), where P is likely to be a limiting nutrient. To investigate the effects of nutrient stoichiometry on planktonic and sediment ecosystem components and processes, we conducted a replicated in situ mesocosm experiment in Lagunita, a shallow pond located in the southwest region of the basin. Inorganic N and P were periodically added to mesocosms under three different N:P regimes (P only, N:P = 16 and N:P = 75) while the control mesocosms were left unamended. After three weeks of fertilization, more than two thirds of the applied P was immobilized into seston or sediment. The rapid uptake of P significantly decreased biomass C:P and N:P ratios, supporting the hypothesis that Lagunita is P-limited. Meanwhile, simultaneous N and P enrichment significantly enhanced planktonic growth, increasing total planktonic biomass by more than 2-fold compared to the unenriched control. With up to 76% of added N sequestered into the seston, it is suspected that the Lagunita microbial community also experienced strong N-limitation. However, when N and P were applied at N:P = 75, the microbes remained in a P-limitation state as in the untreated control. Two weeks after the last fertilizer application, seston C:P and N:P ratios returned to initial levels but chlorophyll a and seston C concentrations remained elevated. Additionally, no P release from the sediment was observed in the fertilized mesocosms. Overall, this study provides evidence that Lagunita is highly sensitive to nutrient perturbation because the biota is primarily P-limited and experiences a secondary N-limitation despite its high TN:TP ratio. This study serves as a strong basis to justify the need for protection of CCB ecosystems and other low-nutrient microbe-dominated systems from anthropogenic inputs of both N and P.

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