, Volume 10, Issue 7, pp 1133–1147 | Cite as

Changes in Canopy Processes Following Whole-Forest Canopy Nitrogen Fertilization of a Mature Spruce-Hemlock Forest

  • E. Gaige
  • D. B. Dail
  • D. Y. Hollinger
  • E. A. Davidson
  • I. J. Fernandez
  • H. Sievering
  • A. White
  • W. Halteman


Most experimental additions of nitrogen to forest ecosystems apply the N to the forest floor, bypassing important processes taking place in the canopy, including canopy retention of N and/or conversion of N from one form to another. To quantify these processes, we carried out a large-scale experiment and determined the fate of nitrogen applied directly to a mature coniferous forest canopy in central Maine (18–20 kg N ha−1 y−1 as NH4NO3 applied as a mist using a helicopter). In 2003 and 2004 we measured NO3 , NH4 +, and total dissolved N (TDN) in canopy throughfall (TF) and stemflow (SF) events after each of two growing season applications. Dissolved organic N (DON) was greater than 80% of the TDN under ambient inputs; however NO3 accounted for more than 50% of TF N in the treated plots, followed by NH4 + (35%) and DON (15%). Although NO3 was slightly more efficiently retained by the canopy under ambient inputs, canopy retention of NH4 +as a percent of inputs increased markedly under fertilization. Recovery of less than 30% of the fertilizer N in TF suggested that the forest canopy retained more than 70% of the applied N (>80% when corrected for N which bypassed tree surfaces at the time of fertilizer addition). Results from plots receiving 15N enriched NO3 and NH4 + confirmed bulk N estimations that more NO3 than NH4 + was washed from the canopy by wet deposition. The isotope data did not show evidence of canopy nitrification, as has been reported in other spruce forests receiving much higher N inputs. Conversions of fertilizer-N to DON were observed in TF for both 15NH4 + and 15NO3 additions, and occurred within days of the application. Subsequent rain events were not significantly enriched in 15N, suggesting that canopy DON formation was a rapid process related to recent N inputs to the canopy. We speculate that DON may arise from lichen and/or microbial N cycling rather than assimilation and re-release by tree tissues in this forest. Canopy retention of experimentally added N may meet and exceed calculated annual forest tree demand, although we do not know what fraction of retained N was actually physiologically assimilated by the plants. The observed retention and transformation of DIN within the canopy demonstrate that the fate and ecosystem consequences of N inputs from atmospheric deposition are likely influenced by forest canopy processes, which should be considered in N addition studies.


nitrogen deposition canopy fertilization canopy N retention throughfall stemflow 15N tracer 



We thank the International Paper Company, Ltd., and GMO, LLC for providing access to the research site in Howland, Maine. We also thank our technical support staff; Mr. John Lee, Ms. Holly Hughes and Mr. Charles Rodrigues and undergraduates; Mr. David Dunston, Ms. Nell Burger and Ms. Gretchen Miles; Department of Energy SURE program interns who worked on this project. This research was supported by the Office of Science (BER), U.S. Department of Energy, through the Northeast Regional Center of the National Institute for Global Environmental Change under Cooperative Agreement Nos. DE-FC02-03ER63613, DE-FC-03-90ER61010, and DE-FG02-00ER63002; DOE-NICCR agreement No, DE-FG02-06ER64157 and NSF award No. 0223188. Financial support does not constitute an endorsement by DOE or NSF of the views expressed in this article. We also thank several anonymous reviewers. This is Maine Agriculture and Forestry Experiment Station publication #2964.

Supplementary material

10021_2007_9081_Fig4_ESM.jpg (1 mb)

Areal spraying of Howland Forest: Dissolved ammonium nitrate (18 kg N/ha/growing season) was applied to the canopy in 5 equally spaced doses from May to September.


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Copyright information

© Springer Science+Business Media, LLC 2007

Authors and Affiliations

  • E. Gaige
    • 1
  • D. B. Dail
    • 1
  • D. Y. Hollinger
    • 2
  • E. A. Davidson
    • 3
  • I. J. Fernandez
    • 1
  • H. Sievering
    • 4
  • A. White
    • 5
  • W. Halteman
    • 6
  1. 1.Department of Plant, Soil, and Environmental SciencesUniversity of MaineOronoUSA
  2. 2.US Forest ServiceNorthern Region Research StationDurhamUSA
  3. 3.Woods Hole Research CenterFalmouthUSA
  4. 4.Geography and Environmental SciencesUniversity of Colorado-DenverDenverUSA
  5. 5.Forest Ecosystem ScienceSchool of Forest Resources, University of MaineOronoUSA
  6. 6.Math and StatisticsUniversity of MaineOronoUSA

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