Needle-age related variability in nitrogen, mobile carbohydrates, and δ13C within Pinus koraiensis tree crowns

Abstract

For both ecologists and physiologists, foliar physioecology as a function of spatially and temporally variable environmental factors such as sunlight exposure within a tree crown is important for understanding whole tree physiology and for predicting ecosystem carbon balance and productivity. Hence, we studied concentrations of nitrogen (N), non-structural carbohydrates (NSC = soluble sugars + starch), and δ(13)C in different-aged needles within Pinus koraiensis tree crowns, to understand the needle age- and crown position-related physiology, in order to test the hypothesis that concentrations of N, NSC, and δ(13)C are needle-age and crown position dependent (more light, more photosynthesis affecting N, NSC, and δ(13)C), and to develop an accurate sampling strategy. The present study indicated that the 1-yr-old needles had significantly higher concentration levels of mobile carbohydrates (both on a mass and an area basis) and N(area) (on an area basis), as well as NSC-N ratios, but significantly lower levels of N(mass) (on a mass basis) concentration and specific leaf area (SLA), compared to the current-year needles. Azimuthal (south-facing vs. north-facing crown side) effects were found to be significant on starch [both on a mass (ST(mass)) and an area basis (ST(area))], δ(13)C values, and N(area), with higher levels in needles on the S-facing crown side than the N-facing crown side. Needle N(mass) concentrations significantly decreased but needle ST(mass), ST(area), and δ(13)C values significantly increased with increasing vertical crown levels. Our results suggest that the sun-exposed crown position related to photosynthetic activity and water availability affects starch accumulation and carbon isotope discrimination. Needle age associated with physiological activity plays an important role in determining carbon and nitrogen physiology. The present study indicates that across-scale sampling needs to carefully select tissue samples with equal age from a comparable crown position.