Vertical gradients and tree-to-tree variation in shoot morphology and foliar nitrogen in an old-growth Pinus strobus stand

Abstract

Acclimation to ambient light has been assumed to be the principal cause of vertical gradients in leaf nitrogen, foliar morphology, and related traits in forest canopies. We examined the relative role of crown exposure, damage, and reproduction as correlates of vertical gradients in shoot and needle morphology, anatomy, and chemistry in a ~120-year-old stand of Pinus strobus L. (eastern white pine) in central Ontario. Internodes at the top of trees were longer, wider, and produced more fascicles, but had lower fascicle survivorship than lower-canopy shoots. Needles on upper-canopy shoots also had higher allocation to resin ducts than mid- or lower-canopy needles, contained less nitrogen on a mass basis, and showed a higher leaf mass per area, C/N ratio, and chlorophyll a/b ratio. Tree-to-tree variation among the 26 trees measured traits was high, owing, in part, to differences in crown damage and reproductive status. Crown damage was associated with reduced leaf mass per area and other traits associated with shade acclimation, while high cone production was associated with reduced leaf nitrogen and chlorophyll in the upper canopy. Our results suggest that factors other than light acclimation play an important role in determining vertical gradients in foliar morphology, nitrogen, and leaf production in forest canopies.