The elemental stoichiometry of aquatic and terrestrial ecosystems and its relationships with organismic lifestyle and ecosystem structure and function: a review and perspectives

Abstract

C, N and P are three of the most important elements used to build living beings, and their uptake from the environment is consequently essential for all organisms. We have reviewed the available studies on water, soils and organism elemental content ratios (stoichiometry) with the aim of identifying the general links between stoichiometry and the structure and function of organisms and ecosystems, in both aquatic and terrestrial contexts. Oceans have variable C:N:P ratios in coastal areas and a narrow range approximating the Redfield ratio in deep water and inner oceanic areas. Terrestrial ecosystems have a general trend towards an increase in soil and plant N:P ratios from cool and temperate to tropical ecosystems, but with great variation within each climatic area. The C:N:P content ratio (from now on C:N:P ratio) is more constrained in organisms than in the water and soil environments they inhabit. The capacity to adjust this ratio involves several mechanisms, from leaf re-absorption in plants to the control of excretion in animals. Several differences in C:N:P ratios are observed when comparing different taxa and ecosystems. For freshwater ecosystems, the growth rate hypothesis (GRH), which has consistent experimental support, states that low N:P supply determines trophic web structures by favoring organisms with a high growth rate. For terrestrial organisms, however, evidence not yet conclusive on the relevance of the GRH. Recent studies suggest that the N:P ratio could play a role, even in the evolution of the genomes of organisms. Further research is warranted to study the stoichiometry of different trophic levels under different C:N:P environment ratios in long-term ecosystem-scale studies. Other nutrients such as K or Fe should also be taken into account. Further assessment of the GRH requires more studies on the effects of C:N:P ratios on anabolic (growth), catabolic (respiration), storage and/or defensive allocation. Combining elemental stoichiometry with metabolomics and/or genomics should improve our understanding of the coupling of different levels of biological organization, from elemental composition to the structure and evolution of ecosystems, via cellular metabolism and nutrient cycling.