Leaf structural and photosynthetic characteristics, and biomass allocation to foliage in relation to foliar nitrogen content and tree size in three Betula species

Young trees 0.03-1.7 m high of three coexisting Betula species were investigated in four sites of varying soil fertility, but all in full daylight, to separate nutrient and plant size controls on leaf dry mass per unit area (MA), light-saturated foliar photosynthetic electron transport rate (J) and the fraction of plant biomass in foliage (FL). Because the site effect was generally non-significant in the analyses of variance with foliar nitrogen content per unit dry mass (NM) as a covariate, NM was used as an explaining variable of leaf structural and physiological characteristics. Average leaf area (S) and dry mass per leaf scaled positively with NM and total tree height (H) in all species. Leaf dry mass per unit area also increased with increasing H, but decreased with increasing NM, whereas the effects were species-specific. Increases in plant size led to a lower and increases in NM to a greater FL and total plant foliar area per unit plant biomass (LAR). Thus, the self-shading probably increased with increasing NM and decreased with increasing H. Nevertheless, the whole-plant average MA, as well as MA values of topmost fully exposed leaves, correlated with NM and H in a similar manner, indicating that scaling of MA with NM and H did not necessarily result from the modified degree of within-plant shading. The rate of photosynthetic electron transport per unit dry mass (JM) scaled positively with NM, but decreased with increasing Hand MA. Thus, increases in MA with tree height and decreasing nitrogen content not only resulted in a lower plant foliar area (LAR = FL/MA), but also led to lower physiological activity of unit foliar biomass. The leaf parameters (JM, NM and MA) varied threefold, but the whole-plant characteristic FL varied 20-fold and LAR 30-fold, indicating that the biomass allocation was more plastically adjusted to different plant internal nitrogen contents and to tree height than the foliar variables. Our results demonstrate that: (1) tree height and NM may independently control foliar structure and physiology, and have an even greater impact on biomass allocation; and (2) the modified within-plant light availabilities alone do not explain the observed patterns. Although there were interspecific differences with respect to the statistical significance of the relationships, all species generally fit common regressions. However, these differences were consistent, and suggested that more competitive species with inherently larger growth rates also more plastically respond to N and H.