Interacting effects of soil nitrogen supply and light availability on understory sapling growth and foliar attributes

Abstract

Light availability in forest understories is a well recognized constraint on sapling growth, but limitations in soil nitrogen (N) availability, and the link to foliar photosynthetic capacity, typically receive less consideration in describing stand dynamics. My primary hypothesis is that light and soil N availability have speciesspecific effects on photosynthetic activity and growth, and that together these resources will better define understory development in complex forests. To test these relationships, I examined 1) soil N indices and the tradeoffs between soil fertility and light attenuation in old-growth forest understories; 2) the effects of light and N constraints on understory sapling foliar N concentration (N%), N per unit area (Na), and natural abundance of C; 3) the effects of light and soil N supply on species growth and photosynthetic activity in a factorial field experiment; and 4) the mechanisms responsible for the stagnation of understory saplings. Soil N indices incorporating dissolved inorganic N and organic N were useful in characterizing differences in N supply among contrasting sites. Understory light availability declined with increasing soil N supply, while understory Abies lasiocarpa had strong correlations between foliar N% and soil N availability, despite shading effects. In partial-cut forests, understory Tsuga heterophylla and Picea glauca x sitchensis had consistent foliar N% across gradients of light availability; in contrast, foliar N% of Betula papyrifera and Thuja plicata declined with increasing shade, which would distort assessments of soil fertility and perhaps contribute to increased mortality of these species in deep shade. Strong correlations between foliar Na and C or growth increment suggest foliar N per unit area is the simplest integration of light availability and N nutrition on leaf photosynthetic activity. Ontogenic interactions that occur among foliar attributes and tree size in forest understories, especially for saplings < 1 m in height, contribute to time effects on growth patterns and emphasize the need for long-term studies of species autecology and stand dynamics. My experimental manipulation of light and N supply on saplings was ineffective, and future research using natural gradients in site productivity may be more fruitful in defining species response to light and N interactions.