Stoichiometric Plasticity Research Articles

Biodiversity and Stoichiometric Plasticity Increase Pico‐Phytoplankton Contributions to Marine Net Primary Productivity and the Biological Pump

AbstractEarth System Models generally predict increasing upper ocean stratification from 21st century warming, which will cause a decrease in the vertical nutrient flux forcing declines in marine net primary productivity (NPP) and carbon export. Recent advances in quantifying marine ecosystem carbon to nutrient stoichiometry have identified large latitudinal and biome variability, with low‐latitude oligotrophic systems harboring pico‐sized phytoplankton exhibiting large phosphorus to carbon cellular plasticity. The climate forced changes in nutrient flux stoichiometry and phytoplankton community composition are thus likely to alter the ocean's biogeochemical response and feedback with the carbon‐climate system. We have added three pico‐phytoplankton functional types within the Biogeochemical Elemental Cycling component of the Community Earth System Model while incorporating variable cellular phosphorus to carbon stoichiometry for all represented phytoplankton types. The model simulates Prochlorococcus and Synechococcus populations that dominate the productivity and sinking carbon export of the tropical and subtropical ocean, and pico‐eukaryote populations that contribute significantly to productivity and export within the subtropical to mid‐latitude transition zone, with the western subtropical regions of each basin supporting the most P‐poor stoichiometries. Subtropical gyre recirculation regions along the poleward flanks of surface western boundary currents are identified as regional hotspots of enhanced carbon export exhibiting C‐rich/P‐poor stoichiometry, preferentially inhabited by pico‐eukaryotes and diatoms. Collectively, pico‐phytoplankton contribute ∼58% of global NPP and ∼46% of global particulate organic carbon export below 100 m through direct and ecosystem processing pathways. Biodiversity and cellular nutrient plasticity in marine pico‐phytoplankton combine to increase their contributions to ocean productivity and the biological carbon pump.

Potassium limitation of forest productivity – Part 2: CASTANEA-MAESPA-K shows a reduction in photosynthesis rather than a stoichiometric limitation of tissue formation

Abstract. Potassium (K) availability constrains forest productivity. Brazilian eucalypt plantations are a good example of the K limitation of wood production. Here, we built upon a previously described model (CASTANEA-MAESPA-K) and used it to understand whether the simulated decline in C source under K deficiency was sufficient to explain the K limitation of wood productivity in Brazilian eucalypt plantations. We developed allocation schemes for both C and K and included these in CASTANEA-MAESPA-K. Neither direct limitations of the C-sink activity nor direct modifications of the C allocation by K availability were included in the model. Simulation results show that the model was successful in replicating the observed patterns of wood productivity limitation by K deficiency. Simulations also show that the response of net primary productivity (NPP) is not linear with increasing K fertilisation. Simulated stem carbon use and water use efficiencies decreased with decreasing levels of K availability. Simulating a direct stoichiometric limitation of NPP or wood growth was not necessary to reproduce the observed decline of productivity under K limitation, suggesting that K stoichiometric plasticity could be different to that of N and P. Confirming previous results from the literature, the model simulated an intense recirculation of K in the trees, suggesting that retranslocation processes were essential for tree functioning. Optimal K fertilisation levels calculated by the model were similar to nutritional recommendations currently applied in Brazilian eucalypt plantations, paving the way for validation of the model at a larger scale and of this approach for developing decision-making tools to improve fertilisation practices.

Interactive effects of warming and eutrophication on zooplankton could reverse the stoichiometric mismatch with phytoplankton

The core ecosystem functioning (e.g. trophic transfer efficiency) is at risk of being disrupted by the growing mismatch between nutrient content of primary producers and nutrient demand of grazing consumers. Ecological stoichiometry provides a conceptual framework that explains this trophic interaction using C, N and P elemental composition across trophic levels. In light of ongoing climate change and eutrophication, previous studies have raised concerns regarding the growing stoichiometric mismatch between phytoplankton and zooplankton, given the stoichiometric plasticity of phytoplankton. However, there is currently little conclusive evidence on the stoichiometric mismatch from a dual perspective of phytoplankton and zooplankton. To address this, we conducted a mesocosm experiment to investigate the separate and combined effects of climate warming (a constant increase of +3.5 ​°C plus heat waves) and eutrophication (nutrient addition) on stoichiometric mismatch between phytoplankton and zooplankton by examining stoichiometric changes in both communities. We observed a growing trend in stoichiometric mismatches when warming or nutrient addition acted individually, which was mediated by the increase in nutrient demand (N, P elements) of zooplankton growth. However, when these stressors acted jointly, the mismatches were reversed. This could be because climate warming and eutrophication combined would lead to changes in species composition, which accordingly reshaped the stoichiometric composition at the community level. These results illustrate the need of stoichiometric mismatches for understanding the implication of global change on trophic interactions and ecosystem functioning, requiring consideration not only of cross-trophic levels but also of compositional changes within communities.

Food sources drive temporal variation in elemental stoichiometry of benthic consumers

AbstractBenthic fauna plays an important role in mediating biogeochemical cycles in coastal areas by storing carbon (C), nitrogen (N), and phosphorus (P) in their body tissues at theoretically homeostatic rates. To maintain homeostasis, the benthic consumers need to be in balance with their resource supply or alter their stoichiometric traits in response to environmental change. However, we lack an understanding regarding the potential variation in the C : N : P content ratios of benthic consumers, especially in marine coastal areas where stoichiometric shifts in macrofauna could have strong effects on sediment carbon and nutrient cycling. By monitoring two sites over a year, we quantified the magnitude and temporal stability of benthic faunal carbon and nutrient pools in coastal soft‐sediment habitats. Our results show that benthic fauna is not strictly homeostatic, but instead expresses temporal variation in C : N : P content. These aquatic consumers undergo ontogenetic changes in diet and morphology, which alter their stoichiometric characteristics. In addition, the faunal C : N : P ratios showed strong seasonal variation at both species and community level, and our results suggest that the stoichiometric traits of benthic consumers shift in response to food sources and environmental conditions. The ability to adapt to varying stoichiometric conditions is essential in face of the growing C : N : P imbalance occurring in marine and coastal ecosystems as a consequence of anthropogenic activities. Therefore, it is critically important to identify the stoichiometric plasticity of different species, before environmental change causes a shift in benthic community composition that will alter functions on an ecosystem level.

Environmental control of marine phytoplankton stoichiometry in the North Atlantic Ocean

The stoichiometric coupling of carbon to limiting nutrients in marine phytoplankton regulates the magnitude of biological carbon sequestration in the ocean. While clear links between plankton C:N ratios and environmental drivers have been identified, the nature and direction of these links, as well as their underlying physiological and ecological controls, remain uncertain. We show, with a well-constrained mechanistic model of plankton ecophysiology, that while nitrogen availability and temperature emerge as the main drivers of phytoplankton C:N stoichiometry in the North Atlantic, the biological mechanisms involved vary depending on the spatiotemporal scale and region considered. We find that phytoplankton C:N stoichiometry is overall controlled by nitrogen availability below 40° N, predominantly driven by ecoevolutionary shifts in the functional composition of the phytoplankton communities, while phytoplankton stoichiometric plasticity in response to dropping temperatures and increased grazing pressure dominates at higher latitudes. Our findings highlight the potential of "organisms-to-ecosystems" modeling approaches based on mechanistic models of plankton biology accounting for physiology, ecology, and trait evolution to explore and explain complex observational data and ultimately improve the predictions of global ocean models.

Greater soil water and nitrogen availability increase C : N ratios of root exudates in a temperate steppe

A better understanding of how rates and stoichiometric signatures of root exudates respond to altered N deposition and rainfall patterns is critical for predicting future impacts of environmental changes on ecosystem function and services. Here, we conducted a field experiment examining the effects of rainfall reduction (−50% compared to the control with normal rainfall), rainfall addition (+50%), N addition (~10 g N m−2 yr−1), and their interactions on root C and N exudation rates and C : N ratios of root exudates from dominant species in a temperate steppe. We hypothesized that increasing soil water and N availability will increase C: N ratios of root exudates because plants will grow more and retain more N in their biomass, thus have lower root N exudation rates. We found greater changes in root N exudation rates (ranging from −18% ~ +11%) than C exudation rates (ranging from −6 ~ +11%) in response to rainfall and N treatments. In addition, N addition and its interactions with rainfall treatments decreased root N exudation rates by 4–18% and therefore increased C : N ratios of root exudates by 8–27% compared to control; however, a contrasting pattern was found under rainfall reduction. Furthermore, changes in root C exudation rates in response to rainfall and N treatments were not related to changes in plant biomass. In contrast, root N exudation rates decreased with increasing plant biomass, resulting in an overall increase in C : N ratios of root exudates with increasing plant biomass. Overall, our results reveal the C : N stoichiometric plasticity of root exudates in response to resource modifications. These findings have important implications for understanding rhizosphere plant-soil-microbe interactions and ecosystem functioning under environmental changes.

Varying influence of phytoplankton biodiversity and stoichiometric plasticity on bulk particulate stoichiometry across ocean basins

Concentrations and elemental ratios of suspended particulate organic matter influence many biogeochemical processes in the ocean, including patterns of phytoplankton nutrient limitation and links between carbon, nitrogen and phosphorus cycles. Here we present direct measurements of cellular nutrient content and stoichiometric ratios for discrete phytoplankton populations spanning broad environmental conditions across several ocean basins. Median cellular carbon-to-phosphorus and nitrogen-to-phosphorus ratios were positively correlated with vertical nitrate-to-phosphate flux for all phytoplankton groups and were consistently higher for cyanobacteria than eukaryotes. Light and temperature were inconsistent predictors of stoichiometric ratios. Across nutrient-rich and phosphorus-stressed biomes in the North Atlantic, but not in the nitrogen-stressed tropical North Pacific, we find that a combination of taxonomic composition and environmental acclimation best predict bulk particulate organic matter composition. Our findings demonstrate the central role of plankton biodiversity and plasticity in controlling linkages between ocean nutrient and carbon cycles in some regions.

Long-term fertilization and residue return affect soil stoichiometry characteristics and labile soil organic matter fractions

Ecological stoichiometry provides the possibility for linking microbial dynamics with soil carbon (C), nitrogen (N), and phosphorus (P) metabolisms in response to agricultural nutrient management. To determine the roles of fertilization and residue return with respect to ecological stoichiometry, we collected soil samples from a 30-year field experiment on residue return (maize straw) at rates of 0, 2.5, and 5.0 Mg ha−1 in combination with 8 fertilization treatments: no fertilizer (F0), N fertilizer, P fertilizer, potassium (K) fertilizer, N and P (NP) fertilizers, N and K (NK) fertilizers, P and K (PK) fertilizers, and N, P, and K (NPK) fertilizers. We measured soil organic C (SOC), total N and P, microbial biomass C, N, and P, water-soluble organic C and N, KMnO4-oxidizable C (KMnO4-C), and carbon management index (CMI). Compared with the control (F0 treatment without residue return), fertilization and residue return significantly increased the KMnO4-C content and CMI. Furthermore, compared with the control, residue return significantly increased the SOC content. Moreover, the NPK treatment with residue return at 5.0 Mg ha−1 significantly enhanced the C:N, C:P, and N:P ratios in the soil, whereas it significantly decreased the C:N and C:P ratios in soil microbial biomass. Therefore, NPK fertilizer application combined with residue return at 5.0 Mg ha−1 could enhance the SOC content through the stoichiometric plasticity of microorganisms. Residue return and fertilization increased the soil C pools by directly modifying the microbial stoichiometry of the biomass that was C limited.

A meta-analysis on environmental drivers of marine phytoplankton C : N : P

Abstract. The elemental stoichiometry of marine phytoplankton plays a critical role in global biogeochemical cycles through its impact on nutrient cycling, secondary production, and carbon export. Although extensive laboratory experiments have been carried out over the years to assess the influence of different environmental drivers on the elemental composition of phytoplankton, a comprehensive quantitative assessment of the processes is still lacking. Here, we synthesized the responses of P:C and N:C ratios of marine phytoplankton to five major drivers (inorganic phosphorus, inorganic nitrogen, inorganic iron, irradiance, and temperature) by a meta-analysis of experimental data across 366 experiments from 104 journal articles. Our results show that the response of these ratios to changes in macronutrients is consistent across all the studies, where the increase in nutrient availability is positively related to changes in P:C and N:C ratios. We found that eukaryotic phytoplankton are more sensitive to the changes in macronutrients compared to prokaryotes, possibly due to their larger cell size and their abilities to regulate their gene expression patterns quickly. The effect of irradiance was significant and constant across all studies, where an increase in irradiance decreased both P:C and N:C. The P:C ratio decreased significantly with warming, but the response to temperature changes was mixed depending on the culture growth mode and the growth phase at the time of harvest. Along with other oceanographic conditions of the subtropical gyres (e.g., low macronutrient availability), the elevated temperature may explain why P:C is consistently low in subtropical oceans. Iron addition did not systematically change either P:C or N:C. Overall, our findings highlight the high stoichiometric plasticity of eukaryotes and the importance of macronutrients in determining P:C and N:C ratios, which both provide us insights on how to understand and model plankton diversity and productivity.

Biological Stoichiometry Regulates Toxin Production in Microcystis aeruginosa (UTEX 2385).

Harmful algal blooms (HABs) are increasing in magnitude, frequency, and duration globally. Even though a limited number of phytoplankton species can be toxic, they are becoming one of the greatest water quality threats to public health and ecosystems due to their intrinsic toxicity to humans and the numerous interacting factors that undermine HAB forecasting. Here, we show that the carbon:nitrogen:phosphorus (C:N:P) stoichiometry of a common toxic phytoplankton species, Microcystis, regulates toxin quotas during blooms through a tradeoff between primary and secondary metabolism. Populations with optimal C:N (< 8) and C:P (< 200) cellular stoichiometry consistently produced more toxins than populations exhibiting stoichiometric plasticity. Phosphorus availability in water exerted a strong control on population biomass and C:P stoichiometry, but N availability exerted a stronger control on toxin quotas by regulating population biomass and C:N:P stoichiometry. Microcystin-LR, like many phytoplankton toxins, is an N-rich secondary metabolite with a C:N stoichiometry that is similar to the optimal growth stoichiometry of Microcystis. Thus, N availability relative to P and light provides a dual regulatory mechanism that controls both biomass production and cellular toxin synthesis. Overall, our results provide a quantitative framework for improving forecasting of toxin production during HABs and compelling support for water quality management that limit both N and P inputs from anthropogenic sources.

Effects of nutrient supply and nutrient ratio on diversity-productivity relationships of phytoplankton in the Cau Hai lagoon, Vietnam.

Diversity and productivity of primary producers are known to be influenced simultaneously by resource availability and resource ratio, but the relative importance of these two factors differed among studies and so far only entire phytoplankton communities were investigated which might ignore specific nutrient requirements and stoichiometric plasticity of different functional groups. We measured nutrient availability (DIN, total N [TN], total P [TP]), nutrient imbalance (TN:TP, DIN:TP, N:Pseston), species richness, and abundance of the whole phytoplankton community, as well as those specific for cyanobacteria, diatoms, and dinoflagellates in Cau Hai lagoon in Vietnam. We determined the correlation among these variables, using structural equation modeling. The models applied to the whole phytoplankton community indicated that the nutrient availability (particularly TP and DIN) drove variation in phytoplankton abundance and richness, and that abundance also depended on species richness. The models applied to different functional groups differed considerably from the entire community and among each other, and only a part of the models was significant. The relationship between nutrient availability (mainly TP) and abundance was driven by cyanobacteria, and the relationship between nutrient imbalance (only with N:Pseston) and species richness was driven by diatoms. Remarkably, the positive relationship between species richness and abundance, as consistently observed for the whole phytoplankton community, was only observed for one of the three functional groups (diatoms), indicating that resource complementarity occurs particularly among species of different functional groups. Our results emphasized that nutrient availability (TP and to a lesser extent DIN) as well as nutrient imbalance (albeit only with N:Pseston as proxy) were driving factors for the phytoplankton community in the Cau Hai lagoon and hence alterations in both of these factors leading to a shift in phytoplankton species composition and productivity.

Parsing the functional specificity of Siderocalin/Lipocalin 2/NGAL for siderophores and related small-molecule ligands

Siderocalin/Lipocalin 2/Neutrophil Gelatinase Associated Lipocalin/24p3 is an innate immune system protein with bacteriostatic activity, acting by tightly binding and sequestering diverse catecholate and mixed-type ferric siderophores from enteric bacteria and mycobacteria. Bacterial virulence achieved through siderophore modifications, or utilization of alternate siderophores, can be explained by evasion of Siderocalin binding. Siderocalin has also been implicated in a wide variety of disease processes, though often in seemingly contradictory ways, and has been proposed to bind to a broader array of ligands beyond siderophores. Using structural, directed mutational, and binding studies, we have sought to rigorously test, and fully elucidate, the Siderocalin recognition mechanism. Several proposed ligands fail to meet rigorous binding criteria, including the bacterial siderophore pyochelin, the iron-chelating catecholamine hormone norepinephrine, and the bacterial second messenger cyclic diguanylate monophosphate. While possessing a remarkably rigid structure, in principle simplifying analyses of ligand recognition, understanding Scn recognition is complicated by the observed conformational and stoichiometric plasticity, and instability, of its bona fide siderophore ligands. Since the role of Siderocalin at the early host/pathogen interface is to compete for bacterial ferric siderophores, we also analyzed how bacterial siderophore binding proteins and enzymes alternately recognize siderophores that efficiently bind to, or evade, Siderocalin sequestration – including determining the crystal structure of Bacillus cereus YfiY bound to schizokinen. These studies combine to refine the potential physiological functions of Siderocalin by defining its multiplexed recognition mechanism.

Stoichiometric plasticity of microbial communities is similar between litter and soil in a tropical rainforest

Heterotrophic microorganisms are commonly thought to be stoichiometrically homeostatic but their stoichiometric plasticity has rarely been examined, particularly in terrestrial ecosystems. Using a fertilization experiment in a tropical rainforest, we evaluated how variable substrate stoichiometry may influence the stoichiometry of microbial communities in the leaf litter layer and in the underlying soil. C:N:P ratios of the microbial biomass were higher in the organic litter layer than in the underlying mineral soil. Regardless of higher ratios for litter microbial communities, C, N, and P fertilization effects on microbial stoichiometry were strong in both litter and soil, without any fundamental difference in plasticity between these two communities. Overall, N:P ratios were more constrained than C:nutrient ratios for both litter and soil microbial communities, suggesting that stoichiometric plasticity arises because of a decoupling between C and nutrients. Contrary to the simplifying premise of strict homeostasis in microbial decomposers, we conclude that both litter and soil communities can adapt their C:N:P stoichiometry in response to the stoichiometric imbalance of available resources.

Sequential nutrient uptake as a potential mechanism for phytoplankton to maintain high primary productivity and balanced nutrient stoichiometry

Abstract. We hypothesize that phytoplankton have the sequential nutrient uptake strategy to maintain nutrient stoichiometry and high primary productivity in the water column. According to this hypothesis, phytoplankton take up the most limiting nutrient first until depletion, continue to draw down non-limiting nutrients and then take up the most limiting nutrient rapidly when it is available. These processes would result in the variation of ambient nutrient ratios in the water column around the Redfield ratio. We used high-resolution continuous vertical profiles of nutrients, nutrient ratios and on-board ship incubation experiments to test this hypothesis in the Strait of Georgia. At the surface in summer, ambient NO3− was depleted with excess PO43− and SiO4− remaining, and as a result, both N : P and N : Si ratios were low. The two ratios increased to about 10 : 1 and 0. 45 : 1, respectively, at 20 m. Time series of vertical profiles showed that the leftover PO43− continued to be removed, resulting in additional phosphorus storage by phytoplankton. The N : P ratios at the nutricline in vertical profiles responded differently to mixing events. Field incubation of seawater samples also demonstrated the sequential uptake of NO3− (the most limiting nutrient) and then PO43− and SiO4− (the non-limiting nutrients). This sequential uptake strategy allows phytoplankton to acquire additional cellular phosphorus and silicon when they are available and wait for nitrogen to become available through frequent mixing of NO3− (or pulsed regenerated NH4). Thus, phytoplankton are able to maintain high productivity and balance nutrient stoichiometry by taking advantage of vigorous mixing regimes with the capacity of the stoichiometric plasticity. To our knowledge, this is the first study to show the in situ dynamics of continuous vertical profiles of N : P and N : Si ratios, which can provide insight into the in situ dynamics of nutrient stoichiometry in the water column and the inference of the transient status of phytoplankton nutrient stoichiometry in the coastal ocean.

Consumer-driven nutrient dynamics in freshwater ecosystems: from individuals to ecosystems.

The role of animals in modulating nutrient cycling [hereafter, consumer-driven nutrient dynamics (CND)] has been accepted as an important influence on both community structure and ecosystem function in aquatic systems. Yet there is great variability in the influence of CND across species and ecosystems, and the causes of this variation are not well understood. Here, we review and synthesize the mechanisms behind CND in fresh waters. We reviewed 131 articles on CND published between 1973 and 1 June 2015. The rate of new publications in CND has increased from 1.4 papers per year during 1973-2002 to 7.3 per year during 2003-2015. The majority of investigations are in North America with many concentrating on fish. More recent studies have focused on animal-mediated nutrient excretion rates relative to nutrient demand and indirect impacts (e.g. decomposition). We identified several mechanisms that influence CND across levels of biological organization. Factors affecting the stoichiometric plasticity of consumers, including body size, feeding history and ontogeny, play an important role in determining the impact of individual consumers on nutrient dynamics and underlie the stoichiometry of CND across time and space. The abiotic characteristics of an ecosystem affect the net impact of consumers on ecosystem processes by influencing consumer metabolic processes (e.g. consumption and excretion/egestion rates), non-CND supply of nutrients and ecosystem nutrient demand. Furthermore, the transformation and transport of elements by populations and communities of consumers also influences the flow of energy and nutrients across ecosystem boundaries. This review highlights that shifts in community composition or biomass of consumers and eco-evolutionary underpinnings can have strong effects on the functional role of consumers in ecosystem processes, yet these are relatively unexplored aspects of CND. Future research should evaluate the value of using species traits and abiotic conditions to predict and understand the effects of consumers on ecosystem-level nutrient dynamics across temporal and spatial scales. Moreover, new work in CND should strive to integrate knowledge from disparate fields of ecology and environmental science, such as physiology and ecosystem ecology, to develop a comprehensive and mechanistic understanding of the functional role of consumers. Comparative and experimental studies that develop testable hypotheses to challenge the current assumptions of CND, including consumer stoichiometric homeostasis, are needed to assess the significance of CND among species and across freshwater ecosystems.

Microzooplankton Stoichiometric Plasticity Inferred from Modeling Mesocosm Experiments in the Peruvian Upwelling Region

Abstract Oxygen minimum zones (OMZs) are often characterised by nitrogen-to-phosphorus (N:P) ratios far lower than the canonical Redfield ratio. Whereas the importance of variable stoichiometry in phytoplankton has long been recognised, variations in zooplankton stoichiometry have received much less attention. Here we combine observations from two shipboard mesocosm nutrient enrichment experiments with an optimality-based plankton ecosystem model, designed to elucidate the roles of different trophic levels and elemental stoichiometry. Pre-calibrated microzooplankton parameter sets represent foraging strategies of dinoflagellates and ciliates in our model. Our results suggest that remineralisation is largely driven by omnivorous ciliates and dinoflagellates, and highlight the importance of intraguild predation. We hypothesize that microzooplankton respond to changes in food quality in terms of nitrogen-to-carbon (N:C) ratios, rather than nitrogen-to-phosphorus (N:P) ratios, by allowing variations in their phosphorus-to-carbon (P:C) ratio. Our results point towards an important biogeochemical role of flexible microzooplankton stoichiometry.

Diagnosing oceanic nutrient deficiency.

The supply of a range of nutrient elements to surface waters is an important driver of oceanic production and the subsequent linked cycling of the nutrients and carbon. Relative deficiencies of different nutrients with respect to biological requirements, within both surface and internal water masses, can be both a key indicator and driver of the potential for these nutrients to become limiting for the production of new organic material in the upper ocean. The availability of high-quality, full-depth and global-scale datasets on the concentrations of a wide range of both macro- and micro-nutrients produced through the international GEOTRACES programme provides the potential for estimation of multi-element deficiencies at unprecedented scales. Resultant coherent large-scale patterns in diagnosed deficiency can be linked to the interacting physical-chemical-biological processes which drive upper ocean nutrient biogeochemistry. Calculations of ranked deficiencies across multiple elements further highlight important remaining uncertainties in the stoichiometric plasticity of nutrient ratios within oceanic microbial systems and caveats with regards to linkages to upper ocean nutrient limitation.This article is part of the themed issue 'Biological and climatic impacts of ocean trace element chemistry'.

Feeding strategies and elemental composition in Ponto‐Caspian peracaridans from contrasting environments: can stoichiometric plasticity promote invasion success?

Summary1. Ponto‐Caspian peracaridans, and mysids and amphipods in particular, are among the most successful aquatic invaders. However, species differ in the trophic‐status range of ecosystems they can invade while establishment rates and impacts can vary substantially between habitats. There is limited knowledge of the environmental factors and species characteristics that drive such variation in invasion success.2. Here we test how trophic level and body stoichiometry vary among peracaridan species and in relation to body size. The amphipod Pontogammarus robustoides and the mysids Limnomysis benedeni and Paramysis lacustris were investigated in ecosystems differing considerably in productivity and nutrient supply, namely an N‐limited eutrophic lagoon and P‐limited mesotrophic lakes.3. As revealed by stable isotope (15N/14N) analysis, herbivory was inferred to be the main feeding mode of L. benedeni. In contrast, the mysid P. lacustris and the amphipod P. robustoides displayed a higher propensity for predatory feeding at larger body sizes, a pattern that was more pronounced in the eutrophic lagoon than in the mesotrophic lakes.4. Their mean stoichiometric composition (P. robustoides C:N:P 108:20:1, L. benedeni 92:21:1 and P. lacustris 93:22:1) demonstrates that these peracaridans are rich in nutrients, especially nitrogen. They all exhibited the same ontogenetic pattern of reduced stoichiometric regulation during juvenile stages and stricter homoeostasis at older stages.5. The higher P content in juveniles of all peracaridan species from the lagoon indicates higher potential somatic and population growth rates than those in the mesotrophic lakes. Such a difference may explain the substantially faster rates of invader establishment observed in the lagoon in comparison with lakes of low trophy.6. Due to differences in ontogenetic and habitat‐induced variation, the study species differed significantly in stoichiometric variability, which was lowest in L. benedeni and highest in P. robustoides. The ranges of species‐specific variation in stoichiometric ratios corresponded to the trophic (by chlorophyll a) and nutrient stoichiometry (N:P) ranges of lentic waters successfully invaded by these species in Lithuania.7. Stoichiometric plasticity, which should be associated with flexibility of feeding strategy, may enhance the potential of peracaridan species to successfully invade habitats with differing trophy and nutrient supply. The optimal feeding strategy should be omnivory with a propensity for predatory feeding, which can be adjusted with respect to ontogenetic nutrient demands and resource availability. Invading species may have a stronger effect on the local biota in ecosystems with high P levels, which promote growth, and N limitation that should favour predation.

Predation risk, stoichiometric plasticity and ecosystem elemental cycling

It is widely held that herbivore growth and production is limited by dietary nitrogen (N) that in turn constrains ecosystem elemental cycling. Yet, emerging evidence suggests that this conception of limitation may be incomplete, because chronic predation risk heightens herbivore metabolic rate and shifts demand from N-rich proteins to soluble carbohydrate-carbon (C). Because soluble C can be limiting, predation risk may cause ecosystem elemental cycling rates and stoichiometric balance to depend on herbivore physiological plasticity. We report on a stoichiometrically explicit ecosystem model that investigates this problem. The model tracks N, and soluble and recalcitrant C through ecosystem compartments. We evaluate how soluble plant C influences C and N stocks and flows in the presence and absence of predation risk. Without risk, herbivores are limited by N and respire excess C so that plant-soluble C has small effects only on elemental stocks and flows. With predation risk, herbivores are limited by soluble C and release excess N, so plant-soluble C critically influences ecosystem elemental stocks flows. Our results emphasize that expressing ecosystem stoichiometric balance using customary C:N ratios that do not distinguish between soluble and recalcitrant C may not adequately describe limitations on elemental cycling.

Intraspecific selectivity, compensatory feeding and flexible homeostasis in the phagotrophic flagellate Oxyrrhis marina: three ways to handle food quality fluctuations

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.