More than colorful: phosphorus allocation to major chemical fractions shifts during leaf development in species exhibiting delayed greening

The colorful "delayed leaf greening" is a common but overlooked phenomenon in phosphorus (P)-limited environments in habitats in tropical, subtropical and temperate forests, but the physiological mechanism underpinning it remains unclear. It is important to understand how allocation of phosphorus to major leaf P fractions shifts during leaf development, as a strategy for utilizing P efficiently. We measured concentrations of leaf nitrogen and P and five chemical P fractions, and eight leaf chemical element concentrations (K, Ca, Mg, Mn, Fe, Cu, Zn and Se) in young and mature leaves of six woody plants exhibiting delayed greening in China. We also measured leaf mass per area, photosynthetic rate, photosynthetic phosphorus-use efficiency, and soil nutrient concentrations. The results indicate six species exhibiting delayed greening had different leaf P concentrations during leaf development, but the same nitrogen concentrations. We further show major leaf chemical P fractions like metabolite P, nucleic acid P and lipid P showed differences in young and mature leaves. The concentration of lipid P, nucleic acid P, Pi and residual P significantly decreased from young to mature leaves, while that of metabolite P was constant. There was a greater allocation of P to phospholipids and metabolite P in mature leaves. The concentration of Cu and K were significantly higher in young leaves. This study provides new insight to investigate the roles of different P fractions in young and mature leaves, and how the allocation shifts for plants to utilize phosphorus.