Organismal Stoichiometry Research Articles

From Elements to Function: Toward Unifying Ecological Stoichiometry and Trait-Based Ecology

The theories developed in ecological stoichiometry are fundamentally based on traits. On the one hand, traits directly linked to cell/body stoichiometry, such as nutrient uptake and storage traits, as well as the associated trade-offs, have the potential to shape ecological interactions such as competition and predation within ecosystems. On the other hand, 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 seldom explicitly integrated in the framework of ecological stoichiometry and, currently, the major challenge is to connect ecological stoichiometry traits with functional traits considered in trait-based ecology. Here, we therefore explore and synthesize existing insights, to develop novel connections between ecological stoichiometry and trait-based ecology. By reviewing key traits and their elemental requirements and illustrating community and ecosystem consequences of variation in elemental balances of traits, we show that unifying the framework of ecological stoichiometry with trait-based ecology sheds light on the interplay between elements and functional traits. Linking elements with functions furthers our understanding of the complex connections between subcellular processes, species interactions, and ultimately ecosystem dynamics.

Stoichiometric traits of stickleback: Effects of genetic background, rearing environment, and ontogeny.

Phenotypes can both evolve in response to, and affect, ecosystem change, but few examples of diverging ecosystem‐effect traits have been investigated. Bony armor traits of fish are good candidates for this because they evolve rapidly in some freshwater fish populations, and bone is phosphorus rich and likely to affect nutrient recycling in aquatic ecosystems. Here, we explore how ontogeny, rearing environment, and bone allocation among body parts affect the stoichiometric phenotype (i.e., stoichiometric composition of bodies and excretion) of threespine stickleback. We use two populations from distinct freshwater lineages with contrasting lateral plating phenotypes (full vs. low plating) and their hybrids, which are mostly fully plated. We found that ontogeny, rearing environment, and body condition were the most important predictors of organismal stoichiometry. Although elemental composition was similar between both populations and their hybrids, we found significant divergence in phosphorus allocation among body parts and in phosphorus excretion rates. Overall, body armor differences did not explain variation in whole body phosphorus, phosphorus allocation, or phosphorus excretion. Evolutionary divergence between these lineages in both allocation and excretion is likely to have important direct consequences for ecosystems, but may be mediated by evolution of multiple morphological or physiological traits beyond plating phenotype.

Investment in boney defensive traits alters organismal stoichiometry and excretion in fish.

Understanding how trait diversification alters ecosystem processes is an important goal for ecological and evolutionary studies. Ecological stoichiometry provides a framework for predicting how traits affect ecosystem function. The growth rate hypothesis of ecological stoichiometry links growth and phosphorus (P) body composition in taxa where nucleic acids are a significant pool of body P. In vertebrates, however, most of the P is bound within bone, and organisms with boney structures can vary in terms of the relative contributions of bones to body composition. Threespine stickleback populations have substantial variation in boney armour plating. Shaped by natural selection, this variation provides a model system to study the links between evolution of bone content, elemental body composition, and P excretion. We measure carbon:nitrogen:P body composition from stickleback populations that vary in armour phenotype. We develop a mechanistic mass-balance model to explore factors affecting P excretion, and measure P excretion from two populations with contrasting armour phenotypes. Completely armoured morphs have higher body %P but excrete more P per unit body mass than other morphs. The model suggests that such differences are driven by phenotypic differences in P intake as well as body %P composition. Our results show that while investment in boney traits alters the elemental composition of vertebrate bodies, excretion rates depend on how acquisition and assimilation traits covary with boney trait investment. These results also provide a stoichiometric hypothesis to explain the repeated loss of boney armour in threespine sticklebacks upon colonizing freshwater ecosystems.

Stoichiometry of Excreta and Excretion Rates of a Stream-dwelling Plethodontid Salamander

Nutrient recycling by fish and amphibians can contribute significantly to ecosystem processes in freshwater ecosystems. Variation in the components that comprise excreta and the factors that control excretion rates may result in differences in the rates and ratios of nutrients excreted by these taxa. These factors can subsequently influence the ecological role of fish and amphibians in freshwater systems. We assessed the composition of excreta of larval Eurycea cirrigera in six headwater streams, and investigated the influence of body size, organismal stoichiometry, and stream nutrient concentrations on mass-specific excretion rates. Excretion components and excretion rates of E. cirrigera were compared to fish in stream and river systems. Excretion of urea comprised the majority of excreted waste in E. cirrigera. This is considerably different from freshwater fish, where the majority of excreted nitrogenous waste is in the form of NH4. Excretion of NH4, soluble reactive phosphorus (SRP), and urea by E. c...

Accumulibacter clades Type I and II performing kinetically different glycogen-accumulating organisms metabolisms for anaerobic substrate uptake

The anaerobic acetate (HAc) uptake stoichiometry of phosphorus-accumulating organisms (PAO) in enhanced biological phosphorus removal (EBPR) systems has been an extensive subject of study due to the highly variable reported stoichiometric values (e.g. anaerobic P-release/HAc-uptake ratios ranging from 0.01 up to 0.93 P-mol/C-mol). Often, such differences have been explained by the different applied operating conditions (e.g. pH) or occurrence of glycogen-accumulating organisms (GAO). The present study investigated the ability of biomass highly enriched with specific PAO clades (‘Candidatus Accumulibacter phosphatis’ Clade I and II, hereafter PAO I and PAO II) to adopt a GAO metabolism. Based on long-term experiments, when Poly-P is not stoichiometrically limiting for the anaerobic VFA uptake, PAO I performed the typical PAO metabolism (with a P/HAc ratio of 0.64 P-mol/C-mol); whereas PAO II performed a mixed PAO-GAO metabolism (showing a P/HAc ratio of 0.22 P-mol/C-mol). In short-term batch tests, both PAO I and II gradually shifted their metabolism to a GAO metabolism when the Poly-P content decreased, but the HAc-uptake rate of PAO I was 4 times lower than that of PAO II, indicating that PAO II has a strong competitive advantage over PAO I when Poly-P is stoichiometrically limiting the VFA uptake. Thus, metabolic flexibility of PAO clades as well as their intrinsic differences are additional factors leading to the controversial anaerobic stoichiometry and kinetic rates observed in previous studies. From a practical perspective, the dominant type of PAO prevailing in full-scale EBPR systems may affect the P-release processes for biological or combined biological and chemical P-removal and recovery and consequently the process performance.

Implications of species addition and decline for nutrient dynamics in fresh waters

Abstract: In terrestrial and aquatic ecosystems, organisms directly affect nutrient storage and cycling by sequestering nutrients via growth and remineralizing nutrients via excretion and egestion. Therefore, species introductions and extirpations can profoundly affect nutrient storage and remineralization rates, and present a challenge for conserving ecosystem function in fresh waters. The literature of consumer-driven nutrient dynamics is growing rapidly, but studies of consumer effects on nutrient storage and remineralization across species and among ecosystems are limited. We compared the effects of 3 grazing taxa, nonnative armored catfish in Mexican streams, native mussels in Oklahoma streams, and native tadpoles in Panamanian streams, on nutrient storage and remineralization. We examined interactions among organismal stoichiometry and biomass, nutrient storage, remineralization rates, and ecosystem size across these 3 groups following species decline (tadpoles and mussels) or introduction (armored ...

Intraspecific variability modulates interspecific variability in animal organismal stoichiometry.

Interspecific differences in organismal stoichiometry (OS) have been documented in a wide range of animal taxa and are of significant interest for understanding evolutionary patterns in OS. In contrast, intraspecific variation in animal OS has generally been treated as analytical noise or random variation, even though available data suggest intraspecific variability in OS is widespread. Here, we assess how intraspecific variation in OS affects inferences about interspecific OS differences using two co-occurring Neotropical fishes: Poecilia reticulata and Rivulus hartii. A wide range of OS has been observed within both species and has been attributed to environmental differences among stream systems. We assess the contributions of species identity, stream system, and the interactions between stream and species to variability in N:P, C:P, and C:N. Because predation pressure can impact the foraging ecology and life-history traits of fishes, we compare predictors of OS between communities that include predators, and communities where predators are absent. We find that species identity is the strongest predictor of N:P, while stream or the interaction of stream and species contribute more to the overall variation in C:P and C:N. Interspecific differences in N:P, C:P, and C:N are therefore not consistent among streams. The relative contribution of stream or species to OS qualitatively changes between the two predation communities, but these differences do not have appreciable effects in interspecific patterns. We conclude that although species identity is a significant predictor of OS, intraspecific OS is sometimes sufficient to overwhelm or obfuscate interspecific differences in OS.

Elemental stoichiometry of basal resources and benthic macroinvertebrates along a land use gradient in a Great Basin watershed

The effects of land use on the elemental stoichiometry of aquatic organisms have rarely been studied in semi-arid watersheds. In eight semi-arid sub-watersheds differing in land use, we determined which predictor variable(s) best explains the elemental variability in two basal food resources and benthic macroinvertebrates (BMI). The elemental composition of periphyton and seston was best explained by percentage of urban and agricultural areas, forested land and associated differences in SRP, DOC, and stream water N:P ratios. In contrast, consumer elemental stoichiometry was related to taxonomic identity and feeding mode. Elemental imbalances were higher for collector-gatherer than for scraper and collector-filterer. However, high spatial and temporal variability in the elemental composition of basal food resources obscured clear spatial patterns of imbalances between nutrient-poor upstream and nutrient-rich downstream sites. Results from this study suggest that land use can affect BMI due to alteration in stoichiometry of their food resources. However, taxonomy and allometry must be taken into account to better understand spatial and temporal changes in the elemental composition of BMI. Our results indicate the importance of considering multiple effects to accurately assess land use effects on producer and consumer stoichiometry, particularly the in highly variable Great Basin watersheds.

Macrobenthic assemblage structure and organismal stoichiometry control faunal processing of particulate organic carbon and nitrogen in oxygen minimum zone sediments

Abstract. The Arabian Sea oxygen minimum zone (OMZ) impinges on the western Indian continental margin between 150 and 1500 m, causing gradients in oxygen availability and sediment geochemistry at the sea floor. Oxygen availability and sediment geochemistry are important factors structuring macrofaunal assemblages in marine sediments. However, relationships between macrofaunal assemblage structure and sea-floor carbon and nitrogen cycling are poorly understood. We conducted in situ 13C:15N tracer experiments in the OMZ core (540 m [O2] = 0.35 μmol l–1) and lower OMZ boundary (800–1100 m, [O2] = 2.2–15.0 μmol l–1) to investigate how macrofaunal assemblage structure, affected by different oxygen levels, and C:N coupling influence the fate of particulate organic matter. No macrofauna were present in the OMZ core. Within the OMZ boundary, relatively high abundance and biomass resulted in the highest macrofaunal assimilation of particulate organic carbon (POC) and nitrogen (PON) at the lower oxygen 800 m stations ([O2] = 2.2–2.36 μmol l–1). At these stations the numerically dominant cirratulid polychaetes exhibited greatest POC and PON uptake. By contrast, at the higher oxygen 1100 m station ([O2] = 15.0 μmol l–1) macrofaunal C and N assimilation was lower, with POC assimilation dominated by one large solitary ascidian. Macrofaunal POC and PON assimilation were influenced by changes in oxygen availability, and significantly correlated to differences in macrofaunal assemblage structure between stations. However, macrofaunal feeding responses were ultimately characterised by preferential organic nitrogen assimilation, relative to their internal C:N budgets.

Widespread intraspecific organismal stoichiometry among populations of the Trinidadian guppy

Summary 1. Ecological stoichiometry expresses ecological interactions as the balance between multiple elements. It relates the ecological function of organisms to their elemental composition, or their organismal stoichiometry. Organismal stoichiometry is thought to reflect elemental investments in life history and morphology acting in concert with variability in abiotic or environmental conditions, but the relative contribution of these factors to natural variability in organismal stoichiometry is poorly understood. 2. We assessed the relative contribution of stream identity, predation, body size and sex to the organismal stoichiometry of guppies (Poecilia reticulata) in six streams in Trinidad. In this system, guppy life-history phenotype evolves in response to predation. Guppies adapted to high-predation (HP) pressure grow faster, mature earlier, produce fewer and smaller offspring and eat a higher-quality diet than guppies adapted to low-predation (LP) pressure. This pattern of life-history evolution is repeated in many rivers encompassing a wide range of abiotic conditions. 3. Organismal stoichiometry of guppies was widely variable, spanning up to ∼70% of the range of variability reported across freshwater fish taxa. Streams from where guppies were sampled were the most important predictor of organismal stoichiometry. In many cases, guppy populations from sites within the same stream varied as much as from sites in different streams. 4. Surprisingly, predation regime was a minor predictor of % C, C : P and C : N in female guppies, despite its strong correlation with life-history phenotype and other organismal traits in this species. Body size and sex were not significant predictors of organismal stoichiometry. 5. Guppies from HP sites were more stoichiometrically balanced with their diets than guppies from LP sites. The latter appeared to be more vulnerable to phosphorus limitation than the former, suggesting that dietary specialization associated with guppy life-history phenotype may have stoichiometric consequences that can affect guppy physiology and nutrient recycling. 6. Our findings suggest that local environmental conditions are a stronger predictor of organismal stoichiometry than organismal traits. We recommend that future work should explicitly consider correlations between organismal traits and organismal stoichiometry in the context of environmental heterogeneity.

Environmental and Organismal Predictors of Intraspecific Variation in the Stoichiometry of a Neotropical Freshwater Fish

The elemental composition of animals, or their organismal stoichiometry, is thought to constrain their contribution to nutrient recycling, their interactions with other animals, and their demographic rates. Factors that affect organismal stoichiometry are generally poorly understood, but likely reflect elemental investments in morphological features and life history traits, acting in concert with the environmental availability of elements. We assessed the relative contribution of organismal traits and environmental variability to the stoichiometry of an insectivorous Neotropical stream fish, Rivulus hartii. We characterized the influence of body size, life history phenotype, stage of maturity, and environmental variability on organismal stoichiometry in 6 streams that differ in a broad suite of environmental variables. The elemental composition of R. hartii was variable, and overlapped with the wide range of elemental composition documented across freshwater fish taxa. Average %P composition was ∼3.2%(±0.6), average %N∼10.7%(±0.9), and average %C∼41.7%(±3.1). Streams were the strongest predictor of organismal stoichiometry, and explained up to 18% of the overall variance. This effect appeared to be largely explained by variability in quality of basal resources such as epilithon N∶P and benthic organic matter C∶N, along with variability in invertebrate standing stocks, an important food source for R. hartii. Organismal traits were weak predictors of organismal stoichiometry in this species, explaining when combined up to 7% of the overall variance in stoichiometry. Body size was significantly and positively correlated with %P, and negatively with N∶P, and C∶P, and life history phenotype was significantly correlated with %C, %P, C∶P and C∶N. Our study suggests that spatial variability in elemental availability is more strongly correlated with organismal stoichiometry than organismal traits, and suggests that the stoichiometry of carnivores may not be completely buffered from environmental variability. We discuss the relevance of these findings to ecological stoichiometry theory.

Strong relationship between elemental stoichiometry and metabolome in plants

Shifts in the elemental stoichiometry of organisms in response to their ontogeny and to changing environmental conditions should be related to metabolomic changes because elements operate mostly as parts of molecular compounds. Here we show this relationship in leaves of Erica multiflora throughout their seasonal development and in response to moderate experimental field conditions of drought and warming. The N/P ratio in leaves decreased in the metabolically active growing seasons, coinciding with an increase in the content of primary metabolites. These results support the growth-rate hypothesis that states that rapidly growing organisms present low N/P ratios because of the increase in allocation of P to RNA. The foliar N/K and P/K ratios were lower in summer and in the drought treatment, in accordance with the role of K in osmotic protection, and coincided with the increase of compounds related to the avoidance of water stress. These results provide strong evidence of the relationship between the changes in foliar C/N/P/K stoichiometry and the changes in the leaf's metabolome during plant growth and environmental stress. Thus these results represent a step in understanding the relationships between stoichiometry and an organism's lifestyle.

The C:N:P stoichiometry of organisms and ecosystems in a changing world: A review and perspectives

This study examined the literature in ISI Web of Science to identify the effects that the main drivers of global change have on the nutrient concentrations and C:N:P stoichiometry of organisms and ecosystems, and examined their relationship to changes in ecosystem structure and function. We have conducted a meta-analysis by comparing C:N:P ratios of plants and soils subjected to elevated [CO2] with those subjected to ambient [CO2]. A second meta-analysis compared the C:N:P ratios of plants and soils that received supplemental N to simulate N deposition and those that did not receive supplemental N. On average, an experimental increase in atmospheric [CO2] increased the foliar C:N ratios of C3 grasses, forbs, and woody plants by 22%, but the foliar ratios of C4 grasses were unaffected. This trend may be enhanced in semi-arid areas by the increase in droughts that have been projected for the coming decades which can increase leaf C:N ratios. The available studies show an average 38% increase in foliar C:P ratios in C3 plants in response to elevated atmospheric [CO2], but no significant effects were observed in C4 grasses. Furthermore, studies that examine the effects of elevated atmospheric [CO2] on N:P ratio (on a mass basis) are warranted since its response remains elusive. N deposition increases the N:P ratio in the plants of terrestrial and freshwater ecosystems, and decreases plants and organic soil C:N ratio (25% on average for C3 plants), reducing soil and water N2 fixation capacity and ecosystem species diversity. In contrast, in croplands subjected to intense fertilization, mostly, animal slurries, a reduction in soil N:P ratio can occur because of the greater solubility and loss of N. In the open ocean, there are experimental observations showing an ongoing increase in P-limited areas in response to several of the factors that promote global change, including the increase in atmospheric [CO2] which increases the demand for P, the warming effect that leads to an increase in water column stratification, and increases in the N:P ratio of atmospheric inputs. Depending on the type of plant and the climate where it grows, warming can increase, reduce, or have no effect on foliar C:N ratios. The results suggest that warming and drought can increase C:N and C:P ratios in warm-dry and temperate-dry terrestrial ecosystems, especially, when high temperatures and drought coincide. Advances in this topic are a challenge because changes in stoichiometric ratios can favour different types of species and change ecosystem composition and structure.

Spatial and temporal variation in the ecological stoichiometry of aquatic organisms in an urban catchment

AbstractUrban land use has increased dramatically over the past few decades. Urban streams are distinguished from forested or agricultural streams, in particular, by their more variable and unpredictable hydrologic pattern. The resulting high variability in nutrient loading is likely to alter the elemental composition of primary producers and, ultimately, to change the elemental composition of other foodweb components. Ecological stoichiometry is a useful framework for improving our understanding of the mass balance of multiple key elements in ecosystems. To this end, the C∶N∶P of key foodweb components were measured in Oxley Creek, an urban catchment in southeastern Queensland, Australia. Ten stream reaches were sampled to explore the spatial variation of C∶N∶P of abundant taxonomic groups across the catchment. Four of these sites were sampled weekly (for 8 wk) to examine temporal variation in elemental composition. Our results suggested that spatial and temporal variation in elemental composition of pri...

To be or not to be what you eat: regulation of stoichiometric homeostasis among autotrophs and heterotrophs

Homeostasis of element composition is one of the central concepts of ecological stoichiometry. In this context, homeostasis is the resistance to change of consumer body composition in response to the chemical composition of consumer's food. To simplify theoretical analysis, it has generally been assumed that autotrophs exhibit flexibility in their composition, while heterotrophs are confined to a constant (strictly homeostatic) body composition. Yet, recent studies suggest that heterotrophs are not universally strictly homeostatic. We examined the degree to which autotrophs and heterotrophs regulate stoichiometric homeostasis (P:C, N:C, N:P, or %P and %N). We conducted a quantitative review and meta-analysis using 132 datasets extracted from 57 literature sources which examined the dependence of organismal stoichiometry on resource stoichiometry. Among individual datasets, there was a wide range of responses from strictly homeostatic to non-homeostatic. Even within heterotrophic organisms, varying levels of homeostasis were observed. Comparing the degree of homeostasis between organisms based on large-scale habitat types using meta-analysis indicated some significant differences between groups. For example, aquatic macroinvertebrates were significantly more homeostatic in terms of P:C than terrestrial invertebrates. Our meta-analysis also confirmed that, with regard to N:P, heterotrophs are significantly more homeostatic than autotrophs. Furthermore, our analysis indicated that the homeostasis parameter 1/H, despite being a potentially useful predictive metric, has to be utilized with caution since it oversimplifies some important aspects of the responses of organisms to elemental imbalances. This critical evaluation of stoichiometric homeostasis contributes to a better understanding of many food-web interactions, which are commonly driven by elemental imbalances between consumers and their resources.

To be or not to be what you eat: regulation of stoichiometric homeostasis among autotrophs and heterotrophs

Homeostasis of element composition is one of the central concepts of ecological stoichiometry. In this context, homeostasis is the resistance to change of consumer body composition in response to the chemical composition of consumer's food. To simplify theoretical analysis, it has generally been assumed that autotrophs exhibit flexibility in their composition, while heterotrophs are confined to a constant (strictly homeostatic) body composition. Yet, recent studies suggest that heterotrophs are not universally strictly homeostatic. We examined the degree to which autotrophs and heterotrophs regulate stoichiometric homeostasis (P:C, N:C, N:P, or %P and %N). We conducted a quantitative review and meta-analysis using 132 datasets extracted from 57 literature sources which examined the dependence of organismal stoichiometry on resource stoichiometry. Among individual datasets, there was a wide range of responses from strictly homeostatic to non-homeostatic. Even within heterotrophic organisms, varying levels of homeostasis were observed. Comparing the degree of homeostasis between organisms based on large-scale habitat types using meta-analysis indicated some significant differences between groups. For example, aquatic macroinvertebrates were significantly more homeostatic in terms of P:C than terrestrial invertebrates. Our meta-analysis also confirmed that, with regard to N:P, heterotrophs are significantly more homeostatic than autotrophs. Furthermore, our analysis indicated that the homeostasis parameter 1/H, despite being a potentially useful predictive metric, has to be utilized with caution since it oversimplifies some important aspects of the responses of organisms to elemental imbalances. This critical evaluation of stoichiometric homeostasis contributes to a better understanding of many food-web interactions, which are commonly driven by elemental imbalances between consumers and their resources.

COMPARING RESOURCE PULSES IN AQUATIC AND TERRESTRIAL ECOSYSTEMS

Resource pulses affect productivity and dynamics in a diversity of ecosystems, including islands, forests, streams, and lakes. Terrestrial and aquatic systems differ in food web structure and biogeochemistry; thus they may also differ in their responses to resource pulses. However, there has been a limited attempt to compare responses across ecosystem types. Here, we identify similarities and differences in the causes and consequences of resource pulses in terrestrial and aquatic systems. We propose that different patterns of food web and ecosystem structure in terrestrial and aquatic systems lead to different responses to resource pulses. Two predictions emerge from a comparison of resource pulses in the literature: (1) the bottom-up effects of resource pulses should transmit through aquatic food webs faster because of differences in the growth rates, life history, and stoichiometry of organisms in aquatic vs. terrestrial systems, and (2) the impacts of resource pulses should also persist longer in terrestrial systems because of longer generation times, the long-lived nature of many terrestrial resource pulses, and reduced top-down effects of consumers in terrestrial systems compared to aquatic systems. To examine these predictions, we use a case study of a resource pulse that affects both terrestrial and aquatic systems: the synchronous emergence of periodical cicadas (Magicicada spp.) in eastern North American forests. In general, studies that have examined the effects of periodical cicadas on terrestrial and aquatic systems support the prediction that resource pulses transmit more rapidly in aquatic systems; however, support for the prediction that resource pulse effects persist longer in terrestrial systems is equivocal. We conclude that there is a need to elucidate the indirect effects and long-term implications of resource pulses in both terrestrial and aquatic ecosystems.

A model of flexible uptake of two essential resources

Microbes require multiple essential elements that they acquire from the environment independently. Here we investigate how microbial stoichiometry and uptake rates depend on the conditions in which they grow. We modify a recent model of growth based on a multinutrient extension of the Droop model to allow a trade-off between ability to acquire two essential resources. In a static analysis, we show that the optimal allocation strategy is the one that results in colimitation by both nutrients. We then add a dynamic equation to model the physiological acclimation uptake rates in changing conditions. This dynamic model predicts that the response of organismal stoichiometry to nutrient supply ratio can vary over time. The response of organismal stoichiometry and growth rate to a nutrient pulse depends on the speed at which cells adapt their uptake rates. In a variable environment, very fast or very slow acclimation may be better strategies than intermediate speed acclimation. We suggest experimental tests of the model and avenues for future model development.

Organismal stoichiometry and the adaptive advantage of variable nutrient use and production efficiency in Daphnia

Models that describe the ecological stoichiometry of individuals and populations often include a simplifying assumption that nutrient ratios in consumers are constant. However, several experiments suggest that organisms can alter their internal stoichiometry under nutrient poor conditions. These experiments also suggest that the production efficiency (biomass produced per unit ingested) of limiting nutrients increases when food quality is poor. We explored how these two physiological adaptations affected growth and reproduction in Daphnia under low nutrient levels by developing three different dynamic simulation models of an individual daphnid under stoichiometric constraints. All three models were converted into individual-based population models interacting with a process-based algal population model. Calibration of the three individual models demonstrated that the assumption of constant nutrient ratios was unable to replicate experimental observations of Daphnia growth under nutrient limited conditions. The addition of increasing phosphorous production efficiency improved the Daphnia's ability to survive and grow, but still not within reported ranges. The addition of variable stoichiometry in the form of increasing nutrient use efficiency yielded an individual daphnid model that could be calibrated across reported food quantity/quality gradients. Further, the model accurately simulated the demonstrated ability of Daphnia to overcome the limitation of poor food quality and eventually suppress the algal population in a high-energy, low-nutrient system. The model accurately predicted the timing of the peak of the Daphnia population as well as the equilibrium biomass levels for both populations reported in a recent experiment. Explorations of model behavior further demonstrated that the adaptations under consideration increase total system P cycling which increases total system production.

Are you what you eat? Physiological constraints on organismal stoichiometry in an elementally imbalanced world

The relative supply of energy and elements available to organisms in the environment has strong effects on their physiology, which, in turn, can alter important ecological processes. Here we consider how resource imbalances affect three basic physiological processes common to all organisms: elemental uptake, incorporation, and release. We review recent research that addresses these core issues (uptake, incorporation, and release) as they relate to elemental homeostasis in autotrophs and heterotrophs. Our review shows the importance that organism elemental homeostasis plays in determining the types of physiological processes used to acquire, assemble, store, and release biogenic elements, which are found in widely varying ratios in the environment. Future research should examine the degree to which organisms assess their internal nutritional composition and that of their food sources within a multiple elemental and biochemical context. Also, scientists should explore if and how the stoichiometry of cellular and molecular responses underlying nutrient (elemental and biochemical) acquisition, incorporation, and release depends on the nutritional composition of food resources. These types of queries will further improve our understanding of the physiological processing of primary elements involved in growth, reproduction, and maintenance of organisms.