Optimal leaf life strategies determine V_c,max dynamic during ontogeny

Leaf photosynthetic properties, for example the maximum carboxylation velocity or V_c,max, change with leaf age due to ontogenetic processes. This study introduces an optimal dynamic allocation scheme to model changes in leaf-level photosynthetic capacity as a function of leaf biochemical constraints (costs of synthesis and defence), nitrogen availability and other environmental factors (e.g. light). The model consists of a system of equations describing RuBisCO synthesis and degradation within chloroplasts, defence and ageing at leaf levels, nitrogen transfer and carbon budget at plant levels. Model results show that optimal allocation principles explained RuBisCO dynamics with leaf age. An approximated analytical solution can reproduce the basic pattern of RuBisCO and V_c,max in rice and in two tropical tree species. The model also reveals leaf life complementarities that remained unexplained in previous approaches, as the interplay between V_c,max at maturation, life span and the decline in photosynthetic capacity with age. Furthermore, it explores the role of defence, which is not implemented in current models. This framework covers some of the existing gaps in integrating multiple processes across plant organs (chloroplast, leaf and whole plant) and is a first-step towards representing mechanistically leaf ontogenetic processes into physiological and ecosystem models.