Leaf size modifies support biomass distribution among stems, petioles and mid-ribs in temperate plants

The implications of extensive variation in leaf size for biomass distribution between physiological and support tissues and for overall leaf physiological activity are poorly understood. Here, we tested the hypotheses that increases in leaf size result in enhanced whole-plant support investments, especially in compound-leaved species, and that accumulation of support tissues reduces average leaf nitrogen (N) content per unit dry mass (N(M)), a proxy for photosynthetic capacity. Leaf biomass partitioning among the lamina, mid-rib and petiole, and whole-plant investments in leaf support (within-leaf and stem) were studied in 33 simple-leaved and 11 compound-leaved species. Support investments in mid-ribs and petioles increased with leaf size similarly in simple leaves and leaflets of compound leaves, but the overall support mass fraction within leaves was larger in compound-leaved species as a result of prominent rachises. Within-leaf and within-plant support mass investments were negatively correlated. Therefore, the total plant support fraction was independent of leaf size and lamina dissection. Because of the lower N(M) of support biomass, the difference in N(M) between the entire leaf and the photosynthetic lamina increased with leaf size. We conclude that whole-plant support costs are weakly size-dependent, but accumulation of support structures within the leaf decreases whole-leaf average N(M), potentially reducing the integrated photosynthetic activity of larger leaves.