Multielement stoichiometry in aquatic invertebrates: When growth dilution matters

Element concentrations in organisms can be variable, often causing deviations from otherwise consistent, taxon-specificmultielement stoichiometries. Such variation can have considerable ecological consequences, yet physiological mechanisms remain unclear. We tested the influence of somatic growth dilution (SGD) on multiple element concentrations under different bioenergetic conditions. SGD occurs when rapid individual growth causes a disproportional gain in biomass relative to gain of a specific element. SGD can strongly affect elements in various organisms, but we lack a general framework to unify results across studies and assess its overall importance.We derived the general conditions that trigger SGD from an element accumulation model. We parameterized the model with bioenergetic and element-specific rates summarized from the literature to compare SGD effects on 15 elements (nonessential metals, essential trace elements, macronutrients) in three aquatic invertebrate taxa. For all taxa, we found that SGD (1) occurs to some degree for all 15 elements over realistic ranges of growth and ingestion rates and (2) has the greatest effect on elements with low efflux (excretion) rates, including certain nonessential metals (e.g., MeHg, Po), essential trace elements, and macronutrients (e.g., N, Fe). Thus, SGD can strongly affect concentrations of a spectrum of elements under natural conditions. These results provide a framework for predicting variation in the elemental composition of animals.