Biomass investment in leaf lamina versus lamina support in relation to growth irradiance and leaf size in temperate deciduous trees.

Foliar biomass investment in support and assimilative compartments was studied in four temperate deciduous tree species along a natural light gradient across the canopy. The species ranked according to shade tolerance as Betula pendula Roth. < Populus tremula L. < Fraxinus excelsior L. < Tilia cordata Mill. Long-term light conditions at sampling locations were characterized as seasonal mean integrated quantum flux density (Q(int), mol m(-2) day(-1)) estimated by a method combining hemispherical photography and light measurements with quantum sensors. Leaf morphology was altered by Q(int) in all species. Both lamina and petiole dry mass per lamina area (LMA and PMA, respectively) increased with increasing Q(int). Shade-tolerant species had lower LMA at low Q(int) than shade-intolerant species; however, PMA was not related to shade tolerance. Across species, the ratio of petiole dry mass to lamina dry mass (PMR) varied from 0.07 to 0.21. It was independent of Q(int) in the simple-leaved species, but decreased with increasing Q(int) in the compound-leaved F. excelsior, which also had the largest foliar biomass investment in petioles. Differences in leaf mass and area, ranging over four orders of magnitude, provided an explanation for the interspecific variability in PMR. Species with large leaves also had greater biomass investments in foliar support than species with smaller leaves. This relationship was similar for both simple- and compound-leaved species. There was a negative relationship between PMR and petiole N concentration, suggesting that petioles had greater carbon assimilation rates and paid back a larger fraction of their construction cost in species with low PMR than in species with high PMR. This was probably the result of a negative relationship between PMR and petiole surface to volume ratio. Nevertheless, petioles had lower concentrations of mineral nutrients than laminas. Across species, the ratio of petiole N to lamina N varied from only 3 to 6%, demonstrating that petiole costs are less in terms of nutrients than in terms of total biomass, and that the petiole contribution to carbon assimilation is disproportionately lower than that of the lamina contribution.