The ratio of foliar nitrogen to foliar phosphorus: a determinant of leaf attributes and height in life-forms of subtropical and tropical plant communities

Abstract

In the species-rich overstorey of tropical and subtropical closed-forests (rainforests), a series of life-forms (emergent trees, canopy trees, subcanopy trees, mid-stratum trees and shrubs, interlaced with lianes) of increasing basal area, height and foliage attributes (leaf area, leaf specific weight and internode length) develop in equilibrium with aerodynamic fluxes (frictional, thermal, evaporative ± atmospheric salinity) in the atmosphere as it flows turbulently over and through a plant community. In both closed-forest and open-forest communities in eastern Australia, the translocation of high-energy nitrogen and phosphorus compounds into developing leaves – during the driest season of the year – increases as soil water becomes more available in the climatic gradient from the subhumid to the per-humid zone. Foliage attributes (leaf area and leaf specific weight) of vertical shoots are determined by the rate of input of high-energy compounds into developing shoot apices. Increasing nutrient input in the transpiration stream results in a greater number of leaves (with similar leaf specific weights) on vertical foliage shoots. The leaf area index of the tree is thus enhanced and leads to increased biomass, basal area and height at maturity. In each life-form within a closed-forest, the size of the root system is allometrically related to aboveground attributes. The ability of the root system to explore available nitrogen and phosphorus stored in the surface soil thus determines the attributes of developing foliage shoots in each of these life-forms. Both leaf areas and leaf specific weights decrease from maxima in canopy trees to leaves of subcanopy and mid-stratum trees in the milder climate under the dense structure of per-humid rainforests. In contrast, in the open-structured, subhumid rainforests, although leaf areas decrease in the gradient from canopy to mid-stratum trees – all exposed to direct solar radiation – leaf specific weights increase as temperatures in the boundary layer around growth-apices increase. The production of nitrate ions in soil, exposed to solar radiation in gaps, increases the uptake of nitrogen into leaves of pioneer trees. Larger and thinner leaves, with higher foliar N : P ratios and nitrate reductase activity, result in and enable rapid regeneration of the rainforest.