Tree Physiology Advance Access originally published online on October 23, 2009
Tree Physiology 2009 29(12):1491-1501; doi:10.1093/treephys/tpp090
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Coupling tree-ring 
13C and 15N to test the effect of fertilization on mature Douglas-fir (Pseudotsuga menziesii var. glauca) stands across the Interior northwest, USA
1 Department of Soil Science, University of Wisconsin, 1525 Observatory Drive, Madison, WI 53706-1299, USA
2 Corresponding author (njbalster{at}wisc.edu)
3 Department of Forest Resources, University of Idaho, Moscow, ID 83844-1133, USA
4 Departments of Plant Pathology and Statistics, 1210W. Dayton St., University of Wisconsin, Madison, WI 53706-1685, USA
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Abstract |
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Nitrogen (N) fertilization causes long-term increases in biomass production in many N-limited forests around the world, but the mechanistic basis underlying the increase is often unclear. One possibility, especially in summer-dry climates, is that N fertilization increases the efficiency with which a finite water supply is consumed to support photosynthesis. This increase is achieved by a reduction in the canopy-integrated concentration of internal CO2 and thus discrimination against 13C. We used stable isotopes of carbon (
13C) in tree rings to experimentally test the physiological impact of N fertilization on mature Douglas-fir (Pseudotsuga menziesii Franco var. glauca) stands across the geographic extent of the Intermountain West, USA. The concentration and the stable isotopes of N (15N) in tree rings were also used to assess the presence and activity of fertilizer N. We hypothesized that N fertilization would (i) increase 15N and N concentration of stemwood relative to non-fertilized stands and (ii) increase stemwood 13C as photosynthetic gas exchange responded to the additional N. This experiment included two rates of urea addition, 178 kg ha–1 (low) and 357 kg ha–1 (high), which were applied twice over a 6-year interval bracketed by the 18 years of wood production measured in this study. Foliar N concentrations measured the year after each fertilization treatment suggest that the fertilizer N had been assimilated by the trees (P < 0.001). The N fertilization significantly enriched stemwood 15N by 1.3
at the low fertilization rate and by 2.4 at the high rate (P < 0.001) despite variation in soil N between sites. However, we found no significant effect of the N fertilizer on 13C of the annual rings (P = 0.76). These data lead us to suggest that alternative mechanisms underlie the growth response to fertilizer, i.e., increase in canopy area and shifts in biomass allocation.
Keywords: carbon isotopes, dual-isotope, nitrogen, nitrogen fertilization, nitrogen isotopes, stable isotopes, tree rings
Received November 5, 2008; Accepted September 22, 2009