A general quantitative theory of forest structure and dynamics

We present the first part of a quantitative theory for the structure and dynamics of forests at demographic and resource steady state. The theory uses allometric scaling relations, based on metabolism and biomechanics, to quantify how trees use resources, fill space, and grow. These individual-level traits and properties scale up to predict emergent properties of forest stands, including size-frequency distributions, spacing relations, resource flux rates, and canopy configurations. Two insights emerge from this analysis: (i) The size structure and spatial arrangement of trees in the entire forest are emergent manifestations of the way that functionally invariant xylem elements are bundled together to conduct water and nutrients up from the trunks, through the branches, to the leaves of individual trees. (ii) Geometric and dynamic properties of trees in a forest and branches in trees scale identically, so that the entire forest can be described mathematically and behaves structurally and functionally like a scaled version of the branching networks in the largest tree. This quantitative framework uses a small number of parameters to predict numerous structural and dynamical properties of idealized forests.