Turgor-limited predictions of tree growth, height and metabolic scaling over tree lifespans.

Abstract

Increasing evidence suggests that tree growth is sink-limited by environmental and internal controls rather than by carbon availability. However, the mechanisms underlying sink-limitations are not fully understood and thus not represented in large-scale vegetation models. We develop a simple, analytically solved, mechanistic, turgor-driven growth model (TDGM) and a phloem transport model (PTM) to explore the mechanics of phloem transport and evaluate three hypotheses. First, phloem transport must be explicitly considered to accurately predict turgor distributions and thus growth. Second, turgor-limitations can explain growth-scaling with size (metabolic scaling). Third, turgor can explain realistic growth rates and increments. We show that mechanistic, sink-limited growth schemes based on plant turgor limitations are feasible for large-scale model implementations with minimal computational demands. Our PTM predicted nearly uniform sugar concentrations along the phloem transport path regardless of phloem conductance, stem water potential gradients and the strength of sink-demands contrary to our first hypothesis, suggesting that phloem transport is not limited generally by phloem transport capacity per se but rather by carbon demand for growth and respiration. These results enabled TDGM implementation without explicit coupling to the PTM, further simplifying computation. We test the TDGM by comparing predictions of whole-tree growth rate to well-established observations (site indices) and allometric theory. Our simple TDGM predicts realistic tree heights, growth rates and metabolic scaling over decadal to centurial timescales, suggesting that tree growth is generally sink and turgor limited. Like observed trees, our TDGM captures tree-size- and resource-based deviations from the classical ¾ power-law metabolic scaling for which turgor is responsible.