Magnitude of trait differences among pairs of populations that were from environments with distinct levels of P availability (gray) or not (black), that were grown in the same common garden. Only traits that had 10 pairwise comparisons were included in this analysis: somatic growth rate, acquisition, assimilation, and allocation (AAA) and phosphorus content. Points, error bars and line types as in Figure 1.

Stoichiometric Traits Vary Widely Within Species: A Meta-Analysis of Common Garden Experiments


“Ecological stoichiometry,” a framework that focuses explicitly on the balances and flows of chemical elements within and between organisms and ecosystems, has provided crucial insights into many biological patterns and processes. Despite the proliferation of stoichiometrically-focused studies in recent decades and recognition of the potential for rapid evolution of stoichiometric traits, the prevalence of genetic variation in stoichiometric traits within species remains unclear. We compiled data from 30 published common garden studies of a broad range of taxa (including invertebrates, vertebrates, and autotrophs) to examine how genetic variation influences the acquisition, assimilation, allocation (AAA), composition, and excretion of elements. To quantify the extent of genetic variation for a given trait we calculated the absolute mean response ratio from pairwise comparisons of populations within the same common garden (820 population and 708 genotype comparisons). We observed substantial intraspecific variation of stoichiometric traits across populations and among genotypes; however, the magnitude of variation was greater in AAA traits (effect sizes of 20 and 164% for population and genotype contrasts, respectively) and excretion (effect sizes of 52 and 23%) than in content of carbon (2.1 and 3.1%) and nitrogen (4.5 and 24%). These results suggest that the content of some elements may be evolutionarily constrained relative to AAA traits that determine the processing of these elements, and that a sole focus on elemental content would underestimate the importance of intraspecific genetic variation, particularly within populations. Across many trait types the variation was greater among genotypes within a population than across populations. Finally, we compared pairs of populations from environments with different phosphorus (P) availability to pairs of populations with similar P availability. Genetic variation in the traits measured was similar regardless of the P environment from which genotypes were isolated, suggesting that differences in elemental availability across environments do not necessarily drive enhanced trait divergence. Overall, our results highlight the substantial amount of intraspecific variation in stoichiometric traits and underscore the potential importance of intraspecific variation in driving ecological and evolutionary processes.

Frontiers in Ecology and Evolution


Ecological stoichiometry (ES) is a scientific framework that views living systems as composite chemical reactions which, like all physical processes, are governed by the law of mass balance (Sterner and Elser, 2002; Sterner et al., 2015). In doing so, ES focuses explicitly on the balance and flow of elements and energy within and between organisms and ecosystems. The ES framework has applicability across a wide range of disciplines, from astrobiology to cancer research (Elser, 2003; Elser et al., 2007), and can be applied across multiple levels of biological organization and across taxa. It has also been argued that explicit consideration of an organism’s “stoichiometric traits” (i.e., elemental contents and traits that influence the flux of elements between an organism and its environment) is a natural and convenient way to investigate the complex interplay between ecology and evolution (Kay et al., 2005). This is because elements can be traced through space and time, such that genetic variation in stoichiometric traits can be linked mechanistically to variation in environmental elemental availability (Matthews et al., 2011; Leal et al., 2017b).