1. Root, stem and leaf traits are thought to be functionally coordinated to maximize the efficiency of acquiring and using limited resources. However, evidence is mixed for consistent whole-plant trait coordination among woody plants, and we lack a clear understanding of the adaptive value of root traits along soil resource gradients. If fine roots are the below-ground analogue to leaves, then low specific root length (SRL) and high tissue density should be common on infertile soil. Here, we test the prediction that root, stem and leaf traits and relative growth rate respond in unison with soil fertility gradients. 2. We measured fine root, stem and leaf traits and relative growth rate on individual seedlings of 66 tree species grown in controlled conditions. Our objectives were (i) to determine whether multiple root traits align with growth rate, leaf and stem traits and with each other and (ii) to quantify the relationships between community-weighted mean root traits and two strong soil fertility gradients that differed in spatial extent and community composition. 3. At the species level, fast growth rates were associated with low root and stem tissue density and high specific leaf area. SRL and root diameter were not clearly related to growth rate and loaded on a separate principal component from the plant economic spectrum. 4. At the community level, growth rate was positively related to soil fertility, and root tissue density (RTD) and branching were negatively related to soil fertility. SRL was negatively related and root diameter was positively related to soil fertility on the large-scale gradient that included ectomycorrhizal angiosperms. 5. Synthesis. Root, stem and leaf tissue traits of tree seedlings are coordinated and influence fitness along soil fertility gradients. RTD responds in unison with above-ground traits to soil fertility gradients; however, root traits are multidimensional because SRL is orthogonal to the plant economic spectrum. In contrast to leaves, trees are not constrained in the way they construct fine roots: plants can construct high or low SRL roots of any tissue density. High RTD is the most consistent belowground trait that reflects adaptation to infertile soil.Key-words: determinants of plant community diversity and structure, leaf economic spectrum, plant economic spectrum, root branching, root diameter, root economic spectrum, root tissue density, soil fertility, specific root length, wood economic spectrum P erez-Ramos et al. 2012;Reich 2014;de la Riva et al. 2016). Investment in cheap tissue promotes fast growth, but comes at the cost of a short life span, whereas investment in expensive tissue produces long-lived organs that have slower metabolic rates. The hypothesis that leaves, stems and roots are coordinated is intuitive from both evolutionary and biophysical perspectives: being fast at acquiring and processing resources is only advantageous when all organs are operating

No abstract

The worldwide plant economic spectrum hypothesis predicts that leaf, stem, and root traits are correlated across vascular plant species because carbon gain depends on leaves being adequately supplied with water and nutrients, and because construction of each organ involves a trade‐off between performance and persistence. Despite its logical and intuitive appeal, this hypothesis has received mixed empirical support. If traits within species diverge in their responses to an environmental gradient, then interspecific trait correlations could be weakened when measured in natural ecosystems. To test this prediction, we measured relative growth rates (RGR) and seven functional traits that have been shown to be related to fluxes of water, nutrients, and carbon across 56 functionally diverse tree species on (1) juveniles in a controlled environment, (2) juveniles in forest understories, and (3) mature trees in forests. Leaf, stem, and fine root traits of juveniles grown in a controlled environment were closely correlated with each other, and with RGR. Remarkably, the seven leaf, stem, and fine root tissue traits spanned a single dimension of variation when measured in the controlled environment. Forest‐grown juveniles expressed lower leaf mass per area, but higher wood and fine root tissue density, than greenhouse‐grown juveniles. Traits and growth rates were decoupled in forest‐grown juveniles and mature trees. Our results indicate that constraints exist on the covariation, not just the variation, among vegetative plant organs; however, divergent responses of traits within species to environmental gradients can mask interspecific trait correlations in natural environments. Correlations among organs and relationships between traits and RGR were strong when plants were compared in a standardized environment. Our results may reconcile the discrepancies seen among studies, by showing that if traits and growth rates of species are compared across varied environments, then the interorgan trait correlations observed in controlled conditions can weaken or disappear.