Research review. Components of leaf dry mass per area – thickness and density – alter leaf photosynthetic capacity in reverse directions in woody plants

Abstract

The relationships of foliage assimilation capacity per unit area (P^P_max) with leaf dry mass per unit area (LMA) and nitrogen content per unit area (N_P) differ between species and within species grown in different habitats. To gain a more mechanistic insight into the dependencies of P^P_max on LMA and N_P, this literature study based on 597 species from a wide range of earth biomes with woody vegetation examines the relations between leaf photosynthetic capacity and the components of LMA (leaf density (D, dry mass per volume) and thickness (T)), and also the correlations of D and T with leaf nitrogen content and fractional leaf volumes in different tissues. Across all species, P^P_max varied 12-fold and photosynthetic capacity per unit dry mass (P^m_max) 16-fold, N_P 12-fold, and nitrogen per unit dry mass (N_m) 13-fold, LMA 46-fold, D 13-fold, and T 35-fold, indicating that foliar morphology was more plastic than foliar chemistry and assimilation rates. Although there were strong positive correlations between P^P_max and N_P, and between P^m_max and N_m, leaf structure was a more important determinant of leaf assimilation capacities. P^P_max increased with increasing LMA and T, but was independent of D. By contrast, P^m_max scaled negatively with LMA because of a negative correlation between P^m_max and D, and was poorly related to T. Analysis of leaf nitrogen and tissue composition data indicated that the negative relationship between D and P^m_max resulted from negative correlations between D and N_m, D and volumetric fraction of leaf internal air space, and D and symplasmic leaf fraction. Thus, increases in leaf density bring about (1) decreases in assimilative leaf compounds, and (2) extensive modifications in leaf anatomy that may result in increases in intercellular transfer resistance to CO₂. Collectively, (1) and (2) lead to decreased P^m_max, and also modify P^P_max versus LMA relationships.