Carbon limitation, stem growth rate and the biomechanical cause of Corner's rules.

Background and aims: Corner's rules describe a global spectrum from large-leaved plants with thick, sparingly branched twigs with low-density stem tissues and thick piths to plants with thin, highly branched stems with high-density stem tissues and thin piths. The hypothesis was tested that, if similar crown areas fix similar amounts of carbon regardless of leaf size, then large-leaved species, with their distantly spaced leaves, require higher stem growth rates, lower stem tissue densities and stiffnesses, and therefore thicker twigs.
Methods: Structural equation models were used to test the compatibility of this hypothesis with a dataset on leaf size, shoot tip spacing, stem growth rate and dimensions, and tissue density and mechanics, sampling 55 species drawn from across the angiosperm phylogeny from a morphologically diverse dry tropical community. Key results: Very good fit of structural equation models showed that the causal model is highly congruent with the data. Conclusions: Given similar amounts of carbon to allocate to stem growth, larger-leaved species require greater leaf spacing and therefore greater stem extension rates and longer stems, in turn requiring lower-density, more flexible, stem tissues than small-leaved species. A given stem can have high resistance to bending because it is thick (has high second moment of area I) or because its tissues are stiff (high Young's modulus E), the so-called E-I trade-off. Because of the E-I trade-off, large-leaved species have fast stem growth rates, low stem tissue density and tissue stiffness, and thick twigs with wide piths and thick bark. The agreement between hypothesis and data in structural equation analyses strongly suggests that Corner's rules emerge as the result of selection favouring the avoidance of self-shading in the context of broadly similar rates of carbon fixation per unit crown area across species.