In their seminal paper, Goldman et al. suggested that phytoplankton close to maximum growth rate attains a restricted optimal N : P ratio close to the Redfield ratio of molar N : P = 16. Recently, the presence of such a global attractor for optimal phytoplankton stoichiometry has been questioned in models and empirical analyses. As the chemical composition of phytoplankton is of major importance for our understanding of global elemental cycles and biogeochemical transformations, we assembled 55 data sets of phytoplankton growth rate and biomass N : P ratios in a meta‐analysis testing (1) whether phytoplankton N : P converges at high growth rates, (2) whether N : P ratios scale with growth rate, and (3) whether the optimal N : P ratios achieved at highest growth rates reflect organism traits or environmental conditions. Across systems and species, phytoplankton N : P decreased with increasing growth rate and at the same time showed decreasing variance, i.e., fast‐growing phytoplankton is more P rich and has a more confined elemental composition. Optimal N : P increased with increasing N : P of available nutrients, i.e., with increasing P limitation. Other differences were rare, except cyanobacteria showed higher optimal N : P than diatoms. Understanding the role of phytoplankton in biogeochemical transformation requires modeling approaches that are stoichiometrically flexible to reflect the dynamics of growth and nutrient supply in primary producers.

Herbivores are generally faced with a plethora of resources which differ in quality. Therefore, they should be able to select foods which most closely match their metabolic needs. Here, we tested the hypothesis that copepods of the species Acartia tonsa select prey cells based on quality differences within prey species. We assessed age‐specific variation in feeding behaviour and evaluated the potential consequences of such variation for nutrient cycles. Nauplii (young) stages characterized by a low nitrogen to phosphorus (N:P) ratio in their body tissue selected for phosphorus‐rich food, while older copepodite stages with higher body N:P selected for nitrogen‐rich food. Further, the analysis of a 35‐year data set in the southern North Sea revealed a positive correlation between the abundance of nauplii and the ratio of dissolved inorganic N:P, thus suggesting that P‐availability for primary producers declines with the population densities of nauplii. Our findings demonstrate that a combination of stage‐specific selective feeding and body stoichiometry has the potential to affect cycling of limiting nutrients when consumer populations change in composition.

Herbivory is more prevalent in the tropics than at higher latitudes. If differences in ambient temperature are the direct cause for this phenomenon, then the same pattern should be visible in a seasonal gradient, as well as in experiments manipulating temperature. Using (15)N stable isotope analyses of natural populations of the copepod Temora longicornis we indeed observed seasonal differences in the trophic level of the copepod and a decrease in trophic level with increasing temperature. In a grazing experiment, with a mixed diet of the cryptophyte Rhodomonas salina and the heterotrophic dinoflagellate Oxyrrhis marina, T. longicornis preferred the cryptophyte at higher temperatures, whereas at lower temperatures it preferred the non-autotrophic prey. We explain these results by the higher relative carbon content of primary producers compared to consumers, in combination with the higher demand for metabolic carbon at higher temperatures. Thus, currently increasing temperatures may cause changes in dietary preferences of many consumers.

Mesocosm experiments coupled with dilution grazing experiments were carried out during the phytoplankton spring bloom 2009. The interactions between phytoplankton, microzooplankton and copepods were investigated using natural plankton communities obtained from Helgoland Roads (54°11.3 0 N; 7°54.0 0 E), North Sea. In the absence of mesozooplankton grazers, the microzooplankton rapidly responded to different prey availabilities; this was most pronounced for ciliates such as strombidiids and strobilids. The occurrence of ciliates was strongly dependent on specific prey and abrupt losses in their relative importance with the disappearance of their prey were observed. Thecate and athecate dinoflagellates had a broader food spectrum and slower reaction times compared with ciliates. In general, high microzooplankton potential grazing impacts with an average consumption of 120% of the phytoplankton production (P p ) were measured. Thus, the decline in phytoplankton biomass could be mainly attributed to an intense grazing by microzooplankton. Copepods were less important phytoplankton grazers consuming on average only 47% of P p . Microzooplankton in turn contributed a substantial part to the copepods' diets especially with decreasing quality of phytoplankton food due to nutrient limitation over the course of the bloom. Copepod grazing rates exceeded microzooplankton growth, suggesting their strong top-down control potential on microzooplankton in the field.Selective grazing by microzooplankton was an important factor for stabilising a bloom of less-preferred diatom species in our mesocosms with specific species (Thalassiosira spp., Rhizosolenia spp. and Chaetoceros spp.) dominating the bloom. This study demonstrates the importance of microzooplankton grazers for structuring and controlling phytoplankton spring blooms in temperate waters and the important role of copepods as top-down regulators of microzooplankton.

Stoichiometric homeostasis is the ability of an organism to keep its body chemical composition constant, despite varying inputs. Stoichiometric homeostasis therefore constrains the metabolic needs of consumers which in turn often feed on resources not matching these requirements. In a broader context, homeostasis also relates to the capacity of an organism to maintain other biological parameters (e.g. body temperature) at a constant level over ambient environmental variations. Unfortunately, there are discrepancies in the literature and ecological and physiological definitions of homeostasis are disparate and partly contradictory. Here, we address this matter by reviewing the existing knowledge considering two distinct groups, regulators and conformers and, based on examples of thermo- and osmoregulation, we propose a new approach to stoichiometric homeostasis, unifying ecological and physiological concepts. We suggest a simple and precise graphical way to identify regulators and conformers: for any given biological parameter (e.g. nutrient stoichiometry, temperature), a sigmoidal relation between internal and external conditions can be observed for conformers while an inverse sigmoidal response is characteristic of regulators. This new definition and method, based on well-studied physiological mechanisms, unifies ecological and physiological approaches and is a useful tool for understanding how organisms are affected by and affect their environment.

Increased reactive nitrogen (Nr ) deposition has raised the amount of N available to organisms and has greatly altered the transfer of energy through food webs, with major consequences for trophic dynamics. The aim of this review was to: (i) clarify the direct and indirect effects of Nr deposition on forest and lake food webs in N-limited biomes, (ii) compare and contrast how aquatic and terrestrial systems respond to increased Nr deposition, and (iii) identify how the nutrient pathways within and between ecosystems change in response to Nr deposition. We present that Nr deposition releases primary producers from N limitation in both forest and lake ecosystems and raises plants' N content which in turn benefits herbivores with high N requirements. Such trophic effects are coupled with a general decrease in biodiversity caused by different N-use efficiencies; slow-growing species with low rates of N turnover are replaced by fast-growing species with high rates of N turnover. In contrast, Nr deposition diminishes below-ground production in forests, due to a range of mechanisms that reduce microbial biomass, and decreases lake benthic productivity by switching herbivore growth from N to phosphorus (P) limitation, and by intensifying P limitation of benthic fish. The flow of nutrients between ecosystems is expected to change with increasing Nr deposition. Due to higher litter production and more intense precipitation, more terrestrial matter will enter lakes. This will benefit bacteria and will in turn boost the microbial food web. Additionally, Nr deposition promotes emergent insects, which subsidize the terrestrial food web as prey for insectivores or by dying and decomposing on land. So far, most studies have examined Nr -deposition effects on the food web base, whereas our review highlights that changes at the base of food webs substantially impact higher trophic levels and therefore food web structure and functioning.

The theories developed in ecological stoichiometry (ES) are fundamentally based on traits. Traits directly linked to cell/body stoichiometry, such as nutrient uptake and storage, as well as the associated trade-offs, have the potential to shape ecological interactions such as competition and predation within ecosystems. Further, traits that indirectly influence and are influenced by nutritional requirements, such as cell/body size and growth rate, are tightly linked to organismal stoichiometry. Despite their physiological and ecological relevance, traits are rarely explicitly integrated in the framework of ES and, currently, the major challenge is to more closely inter-connect ES with trait-based ecology (TBE). Here, we highlight four interconnected nutrient trait groups, i.e., acquisition, body stoichiometry, storage, and excretion, which alter interspecific competition in autotrophs and heterotrophs. We also identify key differences between producer-consumer interactions in aquatic and terrestrial ecosystems. For instance, our synthesis shows that, in contrast to aquatic ecosystems, traits directly influencing herbivore stoichiometry in forested ecosystems should play only a minor role in the cycling of nutrients. We furthermore describe how linking ES and TBE can help predict the ecosystem consequences of global change. The concepts we highlight here allow us to predict that increasing N:P ratios in ecosystems should shift trait dominances in communities toward species with higher optimal N:P ratios and higher P uptake affinity, while decreasing N retention and increasing P storage.

The phagotrophic flagellate Oxyrrhis marina shows a strong stoichiometric plasticity when fed differently grown Rhodomonas salina. We tested whether differently pre-conditioned O. marina displayed selective feeding behaviour from a mixture of nitrogen and phosphorus depleted R. salina. We observed selective feeding of O. marina, always selecting phosphorus rich R. salina independent of the pre-conditioning of the protists. In a second experiment, O. marina was again pre-conditioned either with nitrogen-or phosphorus-depleted R. salina and was refed with either of the differently limited R. salina in single food treatments (not in a mixture). The phagotrophic flagellate displayed compensatory feeding which means that O. marina feed more on the food source which they were not given before. Due to its stoichiometric plasticity, O. marina might handle bad quality food by following the stoichiometry of its prey and additionally by active selective feeding towards P-rich algae to enhance growth. Post-ingestion selection might as well be an important feature which means that ingested elements in excess are quickly excreted and scarce elements are ingested through accelerated food uptake.