Inadequate supply of one or more mineral elements can slow the growth of animal consumers and alter their physiology, life history and behaviour. A key concept for understanding nutrient deficiency in animals is the threshold elemental ratio (TER), at which growth limitation switches from one element to another. We used a stoichiometric model that coupled animal bioenergetics and body elemental composition to estimate TER of carbon and phosphorus (TER(C:P)) for 41 aquatic consumer taxa. We found a wide range in TER(C:P) (77-3086, ratio by atoms), which was generated by interspecific differences in body C : P ratios and gross growth efficiencies of C. TER(C:P) also varied among aquatic invertebrates having different feeding strategies, such that detritivores had significantly higher threshold ratios than grazers and predators. The higher TER(C:P) in detritivores resulted not only from lower gross growth efficiencies of carbon but also reflected lower body P content in these consumers. Supporting previous stoichiometric theory, we found TER(C:P) to be negatively correlated with the maximum growth rate of invertebrate consumers. By coupling bioenergetics and stoichiometry, this analysis revealed strong linkages among the physiology, ecology and evolution of nutritional demands for animal growth.

Summary 1. Ecological stoichiometry deals with the mass balance of multiple key elements [e.g. carbon (C), nitrogen (N), phosphorus (P)] in ecological systems. This conceptual framework, largely developed in the pelagic zone of lakes, has been successfully applied to topics ranging from population dynamics to biogeochemical cycling. More recently, an explicit stoichiometric approach has also been used in many other environments, including freshwater benthic ecosystems. 2. Description of elemental patterns among benthic resources and consumers provides a useful starting point for understanding causes of variation and stoichiometric imbalance in feeding interactions. Although there is considerable overlap among categories, terrestrially‐derived resources, such as wood, leaf litter and green leaves have substantially higher C : nutrient ratios than other resources of both terrestrial and aquatic origin, such as periphyton and fine particulate organic matter. The elemental composition of these resources for benthic consumers is modulated by a range of factors and processes, including nutrient availability and ratios, particle size and microbial colonisation. 3. Among consumers in benthic systems, bacteria are the most nutrient‐rich, followed (in descending order) by fishes, invertebrate predators, invertebrate primary consumers, and fungi. Differences in consumer C : nutrient ratios appear to be related to broad‐scale phylogenetic differences which determine body size, growth rate and resource allocation to structural body constituents (e.g. P‐rich bone). 4. Benthic consumers can influence the stoichiometry of dissolved nutrients and basal resources in multiple ways. Direct consumption alters the stoichiometry of food resources by increasing nutrient availability (e.g. reduced boundary layer thickness on substrata) or through removal of nutrient‐rich patches (e.g. selective feeding on fungal patches within leaf litter). In addition, consumers alter the stoichiometry of resources and dissolved nutrient pools through the return of egested or excreted nutrients. In some cases, consumer excretion supplies a large proportion of the nutrients required by algae and heterotrophic microbes and alters elemental ratios of dissolved nutrient pools. 5. Organic matter decomposition in benthic systems is accompanied by significant changes in the elemental composition of organic matter. Microbial colonisation of leaf litter influences C : nutrient ratios, and patterns of microbial succession (e.g. fungi followed by bacteria) may be under some degree of stoichiometric control. Large elemental imbalances exist between particulate organic matter and detritivores, which is likely to constrain growth rates and invertebrate secondary production. Such imbalances may therefore select for behavioural and other strategies for dealing with them. Comminution of large particles by benthic consumers alters detrital C : nutrient ratios and can influence the stoichiometry of elemental export from whole catchments. 6. A stoichiometric framework is l...

2005. Are you what you eat? Physiological constraints on organismal stoichiometry in an elementally imbalanced world. Á/ Oikos 109: 18 Á/28.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.

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We examined how mayfly growth rates and body stoichiometry respond to changing phosphorus (P) content in food. In two experiments, mayfly nymphs were given high or low quantities of food at different carbon:phosphorus (C:P) ratios and their growth was measured. Low food quantity resulted in negative growth rates in both experiments, regardless of food P content. However, under high food availability, mayfly growth was affected by the type of food eaten, with low C:P ratio food producing more rapid growth. In addition, mayfly growth increased somewhat when P‐poor food was artificially enriched with inorganic P although this effect was not statistically significant. Mayfly body P content was inversely related to body size but increased in animals fed artificially P‐enriched food. A model was constructed to simulate mass balance constraints on mayfly growth imposed by the relative supply of two elements (C and P) in food. The model shows that mayfly growth should be limited by food P content at moderately low C:P ratios (c. 120, by mass). Given high C:P ratios (mean c. 270, by mass) in periphyton from oligotrophic boreal lakes, our experimental and theoretical results indicate that stoichiometric constraints are important factors affecting benthic food webs in lakes from the Canadian Shield and perhaps in other systems with similarly high C:P ratios in periphyton.