, Volume 7, Issue 5, pp 440–453 | Cite as

Three-dimensional Structure of an Old-growth Pseudotsuga-Tsuga Canopy and Its Implications for Radiation Balance, Microclimate, and Gas Exchange

  • Geoffrey G. ParkerEmail author
  • Mark E. Harmon
  • Michael A. Lefsky
  • Jiquan Chen
  • Robert Van Pelt
  • Stuart B. Weis
  • Sean C. Thomas
  • William E. Winner
  • David C. Shaw
  • Jerry F. Frankling


We describe the three-dimensional structure of an old-growth Douglas-fir/western hemlock forest in the central Cascades of southern Washington, USA. We concentrate on the vertical distribution of foliage, crowns, external surface area, wood biomass, and several components of canopy volume. In addition, we estimate the spatial variation of some aspects of structure, including the topography of the outer surface, and of microclimate, including the within-canopy transmittance of photosynthetically active radiation (PAR). The crowns of large stems, especially of Douglas-fir, dominate the structure and many aspects of spatial variation. The mean vertical profile of canopy surfaces, estimated by five methods, generally showed a single maximum in the lower to middle third of the canopy, although the height of that maximum varied by method. The stand leaf area index was around 9 m2 m−2, but also varied according to method (from 6.3 to 12.3). Because of the deep narrow crowns and numerous gaps, the outer canopy surface is extremely complex, with a surface area more than 12 times that of the ground below. The large volume included below the outer canopy surface is very porous, with spaces of several qualitatively distinct environments. Our measurements are consistent with emerging concepts about the structure of old-growth forests, where a high degree of complexity is generated by diverse structural features. These structural characteristics have implications for various ecosystem functions. The height and large volume of the stand indicate a large storage component for microclimatic variables. The high biomass influences the dynamics of those variables, retarding rates of change. The complexity of the canopy outer surface influences radiation balance, particularly in reducing short-wave reflectance. The bottom-heaviness of the foliage profile indicates much radiation absorption and gas exchange activity in the lower canopy. The high porosity contributes to flat gradients of most microclimate variables. Most stand respiration occurs within the canopy and is distributed over a broad vertical range.

biomass canopy cover complexity gap hypsograph leaf area index old-growth forests respiration spatial variation transmittance vertical structure 



This work was conducted at the Wind River Canopy Crane Research Facility (WRCCRF), a cooperative scientific venture among the University of Washington, the US Forest Service Pacific Northwest Research Station, and the Gifford Pinchot National Forest. Support was provided by the University of Washington, the National Science Foundation’s LTER program, the Smithsonian Institution, the Global Canopy Programme, NASA’s Goddard Space Flight Center, the Forest Service Pacific Northwest Research Station, and by the Office of Science, US Department of Energy, through the Western Regional Center of the National Institute for Global Environmental Change under cooperative agreement no. DE-FC03-90ER61010. We thank Carrie Bayless, Ken Bible, David Braun, Mark Creighton, David Ford, Annie Hamilton, David Harding, Tom Hinckley, Hiroaki Ishii, Dar Roberts, Mark Rudnicki, Segun Ogunjemiyo, Bo Song, Tom Suchanek, Mark Sumera, Susan Ustin, and Richard Waring for advice, data, discussions, and support.


  1. 1.
    Acker, SA, Halpern, CB, Harmon, ME, Dyrness, CT 2002Trends in bole biomass accumulation, net primary production and tree mortality in Pseudotsuga menziesii forests of contrasting ageTree Physiol222137PubMedGoogle Scholar
  2. 2.
    Baldocchi, DB, Hicks, BB, Meyers, TP 1988Measuring biosphere–atmosphere exchange of biologically related gases with micrometeorological methodsEcology69133140Google Scholar
  3. 3.
    Betts, AK, Ball, JH 1977Albedo over the boreal forestJ Geophys Res10228,90128,909Google Scholar
  4. 4.
    Brockelman, WY 1998Study of tropical forest canopy height and cover using a point-intercept methodDallmeier, FComiskey, JA eds. Forest biodiversity research, monitoring and modelingParthenon and Paris: UNESCOPearl River (NY)52131Google Scholar
  5. 5.
    Campbell, GS, Norman, JM 1998An introduction to environmental biophysicsSpringer-VerlagNew YorkGoogle Scholar
  6. 6.
    Canham, CD, Denslow, JS, Platt, WS, Runkle, WJ,Jr, Spies, TA, White, PS 1990Light regimes beneath closed canopies and tree-fall gaps in temperate and tropical forestsCan J For Res2062031CrossRefGoogle Scholar
  7. 7.
    Cermak, J 1989Solar equivalent leaf area: an efficient biometrical parameter of individual leaves, trees, and standsTree Physiol526989PubMedGoogle Scholar
  8. 8.
    Chen J, Paw U KT, Ustin SL, Suchanek TH, Bond BJ, Brosofske KD, Falk M. 2004. “Net ecosystem exchanges of carbon, water, and energy in young and old-growth Douglas-fir forests.” Ecosystems 7:Google Scholar
  9. 9.
    Cohen, WB, Spies, TA 1992Estimating structural attributes of Douglas-fir/western hemlock forest stands from Landsat and SPOT imageryRemote Sens Environ41117CrossRefGoogle Scholar
  10. 10.
    Cohen, WB, Spies, TA, Bradshaw, GA 1990Semivariograms of digital imagery for analysis of conifer canopy structureRemote Sens Environ3416778CrossRefGoogle Scholar
  11. 11.
    DeAngelis, DL, Gardner, RH, Shugart, HH 1981Productivity of forest ecosystems studied during the IBP: the woodlands data setReichle, DE eds. Dynamic properties of forest ecosystems (UK)Cambridge University PressCambridge567673Google Scholar
  12. 12.
    DeBell, DS, Franklin, JF 1987Old-growth Douglas-fir and western hemlock: a 36-year record of growth and mortalityWest J Appl For21114Google Scholar
  13. 13.
    Dickenson, RE 1983Land surface processes and climate-surface albedos and energy balanceAdv Geophys26305352Google Scholar
  14. 14.
    Dubrasich, ME, Hann, DW, Tappeiner, JC,II 1997Methods for evaluating crown area profiles of forest standsCan J For Res2738592CrossRefGoogle Scholar
  15. 15.
    Ford, ED 1976The canopy of a Scots pine forest: description of a surface of complex roughnessAgric Meteorol17932CrossRefGoogle Scholar
  16. 16.
    Franklin, JF, DeBell, DS 1988Thirty-six years of tree population change in old-growth Pseudotsuga-Tsuga forestCan J For Res186339Google Scholar
  17. 17.
    Franklin JF, Cromack K Jr, Denison W, McKee A, Maser C, Sedell J, Swanson F, and others. 1981. Ecological characteristics of old-growth Douglas-fir forests. USDA Forest Service General Technical Report PNW-118. Portland (OR): Pacific Northwest Research Station.Google Scholar
  18. 18.
    Franklin JF, Spies TA, Van Pelt R, Carey A, Thornburgh D, Burg DR, Lindenmayer D, and others. 2002. Disturbances and the structural development of natural forest ecosystems with some implications for silviculture. For Ecol Manage 155:399–423.Google Scholar
  19. 19.
    Frazer, WG, Trofymow, JA, Lertzman, KP 2000Canopy openness and leaf area in a chronosequence of coastal temperate rainforestsCan J For Res3023956CrossRefGoogle Scholar
  20. 20.
    Harding, DJ, Lefsky, MA, Parker, GG, Blair, B 2001Laser altimeter canopy height profiles: methods and validation for closed-canopy, broadleaf forestsRemote Sens Environ7628397CrossRefGoogle Scholar
  21. 21.
    Harmon ME, Bible K, Shaw D, Remillard S, Sexton J, Fasth B, Priestley A, and others. 1998. Permanent plots surrounding the Wind River canopy crane. Permanent plots of the Pacific Northwest. Report No. 1. http:/www/ Scholar
  22. 22.
    Harmon ME, Bible K, Ryan MG, Shaw D, Chen J, Klopatek J, Li X. 2004. Production, respiration, and overall carbon balance in an old-growth Pseudotuga-Tsuga forest ecosystem. Ecosystems 7:.Google Scholar
  23. 23.
    Hitchcock, CL, Cronquist, A 1978Flora of the Pacific Northwest: an illustrated manualUniversity of Washington PressSeattle (WA)Google Scholar
  24. 24.
    Ishii, H, Reynolds, JH, Ford, ED, Shaw, DC 2000aHeight growth and vertical development of an old-growth Pseudotsuga-Tsuga forest in southwestern Washington State, U.S.ACan J For Res301724CrossRefGoogle Scholar
  25. 25.
    Ishii, H, Clements, JP, Shaw, DC 2000bBranch growth and crown form in old coastal Douglas-firFor Ecol Manage1318191CrossRefGoogle Scholar
  26. 26.
    Isaaks, EH, Srivastava, RM 1989An introduction to applied geostatisticsOxford University PressNew YorkGoogle Scholar
  27. 27.
    Janisch, JE, Harmon, ME 2002Successional changes in live and dead wood carbon stores: implications for net ecosystem productivityTree Physiol227789PubMedGoogle Scholar
  28. 28.
    Jarvis, PG, McNaughton, KG 1986Stomatal control of transpiration: scaling up from leaf to regionAdv Ecol Res15149Google Scholar
  29. 29.
    Jarvis, PG, James, GB, Landsberg, JJ 1976Coniferous forestsMonteith, JJ eds. Vegetation and the atmosphere; vol 2. Case studiesAcademic PressLondon171240Google Scholar
  30. 30.
    Kaimal, JC, Finnigan, JJ 1994Atmospheric boundary layer flows—their structure and measurementOxford University PressNew YorkGoogle Scholar
  31. 31.
    Kuiper LC. 1988. The structure of natural Douglas-fir forests in western Washington and western Oregon. Paper no. 88–5. Wageningen (Netherlands): Agricultural University of Wageningen.Google Scholar
  32. 32.
    Lefsky, MA, Cohen, WB, Acker, SA, Parker, GG, Spies, TA, Harding, D 1999Lidar remote sensing of biophysical properties of canopy structure of forests of Douglas-fir and western hemlockRemote Sens Environ7033961CrossRefGoogle Scholar
  33. 33.
    Lovett, GM, Reiners, WA, Olson, RK 1982Cloud droplet deposition in subalpine balsam fir forests: hydrological and chemical inputsScience21813034Google Scholar
  34. 34.
    Mariscal MJ, Martens SN, Ustin SL, Chen J, Weiss SB, Roberts DA. 2004. Light transmission profiles in an old-growth canopy: simulations of photosynthetically active radiation using spatially explicit radiative transfer models. Ecosystems 7:Google Scholar
  35. 35.
    Marshall, JD, Waring, RH 1986Comparison of methods of estimating leaf-area index of old-growth Douglas-firEcology679759Google Scholar
  36. 36.
    Martens, SN, Ustin, SL, Rousseau, RA 1993Estimation of the canopy leaf area index by gap fraction analysisFor Ecol Manage6191108CrossRefGoogle Scholar
  37. 37.
    Monsi, M, Uchijima, Z, Oikawa, T 1973Structure of foliage canopies and photosynthesisAnnu Rev Ecol Syste430127CrossRefGoogle Scholar
  38. 38.
    Moorcroft, PR, Hurtt, GC, Pacala, SW 2001A method for scaling vegetation dynamics: the Ecosystem Demography model (ED)Ecol Monog7155786Google Scholar
  39. 39.
    Mukammal, EI 1971Some aspects of radiant energy in a pine forestArch Met Geophys Biokl [B]192952Google Scholar
  40. 40.
    Newton T, Paw U KT, Falk M, Shaw RH, King T, Hsiao TC, Pyles RD, and others. 2000. The microclimate of a 65 m tall, old-growth coniferous forest. 24th Conference on Agricultural and Forest Meteorology, Davis, CA, USA, Angnot 2000. American Meteorological SocietyGoogle Scholar
  41. 41.
    Ni, W, Woodcock, CE 2000Effect of canopy structure and the presence of snow on the albedo of boreal conifer forestsJ Geophys Res105(D911,87988Google Scholar
  42. 42.
    Ogunjemiyo S, Parker GG, Roberts D. (Forthcoming) Reflections in bumpy country: implications of canopy surface variations for the energy balance of vegetation. Geosci Remote Sens. Forthcoming.Google Scholar
  43. 43.
    O’Neill, RV, DeAngelis, DL 1970Comparative productivity and biomass relations of forest ecosystemsReichle, DE eds. Dynamic properties of forest ecosystemsCambridge University PressCambridge (UK)41149Google Scholar
  44. 44.
    Parker, GG 1997Canopy structure and light environment of an old-growth Douglas-fir/western hemlock forestNorthwest Sci7126170Google Scholar
  45. 45.
    Parker GG, Russ ME (2004) The canopy surface and stand development: assessing forest canopy structure and complexity with near-surface altimetry. Forest Ecol Manage. 189: 307–315Google Scholar
  46. 46.
    Parker, GG, Lefsky, MA, Harding, DJ 2001Light transmittance in forest canopies determined using airborne laser altimetry and in-canopy quantum measurementsRemote Sens Environ76298309CrossRefGoogle Scholar
  47. 47.
    Parker, GG, Davis, MM, Chapotin, SM 2002Canopy light transmittance in Douglas- fir/western hemlock standsTree Physiol2214757PubMedGoogle Scholar
  48. 48.
    Paw U KT, Falk M, Suchanek TH, Ustin SL, Chen J, Park Y-S, Winner WE, and others. 2004. Carbon dioxide exchange between an old-growth forest and the atmosphere. Ecosystems 7:Google Scholar
  49. 49.
    Peterson, U, Nilson, T 1993Successional reflectance trajectories in northern temperate forestsInt J Remote Sens1460913Google Scholar
  50. 50.
    Pielke, RA, Avissar, R 1990Influence of landscape structure on local and regional climateLandscape Ecol413355Google Scholar
  51. 51.
    Pierce, LL, Running, SW 1988Rapid estimation of coniferous forest leaf area index using a portable integrating radiometerEcology6917627Google Scholar
  52. 52.
    Pike, RJ, Wilson, SE 1971Elevation–relief ratio, hypsometric integral, and geomorphic area–altitude analysisGeol Soc Am Bull82107984Google Scholar
  53. 53.
    Press, WH, Teukolsky, SA, Vetterling, WT, Flannery, BP 1992Numerical recipes in FORTRAN—the art of scientific computing. 2nd edCambridge University PressCambridge (UK)Google Scholar
  54. 54.
    Rich, PM 1990Characterizing plant canopies with hemispherical photographsRemote Sens Environ51329zbMATHGoogle Scholar
  55. 55.
    Roberts DA, Ustin SL, Ogunjemiyo S, Greenberg J, Dobrowski SZ, Chen J, Hinckley TM. 2004. Spectral and structural measures of Northwest forest vegetation at leaf to landscope scales. Ecosytems 7:Google Scholar
  56. 56.
    Shaw DC, Franklin JF, Bible K, Klopatek J, Freeman E, Greene S, Parker GG. 2004. Ecological setting of the Wind River old-growth forests. Ecosystems 7:Google Scholar
  57. 57.
    Smith, NJ 1992Estimating leaf area index and light extinction coefficients in stands of Douglas-fir (Pseudotsuga menziesii)Can J For Res2331721Google Scholar
  58. 58.
    Smithwick, EAH, Harmon, ME, Remillard, SM, Acker, SA, Franklin, JF 2002Potential upper bounds of carbon stores in the Pacific NorthwestEcol Appl12130317Google Scholar
  59. 59.
    Song, B 1998Three-dimensional forest canopies and their spatial relationships to understory vegetation [dissertation]Michigan Technological UniversityHoughton (MI)Google Scholar
  60. 60.
    Spies, T 1998Forest structure: a key to the ecosystemNorthwest Sci72(special issue349CrossRefPubMedGoogle Scholar
  61. 61.
    Spies, TA, Franklin, JF, Klopsch, M 1990Canopy gaps in Douglas-fir forests of the Cascade mountainsCan J For Res206495Google Scholar
  62. 62.
    Stewart, JB 1971The albedo of a pine forestQ J R Meteorol Soc975614CrossRefGoogle Scholar
  63. 63.
    Tajchman, SJ 1972The radiation and energy balances of coniferous and deciduous forestsJ Appl Ecol935975Google Scholar
  64. 64.
    Thomas, SC, Winner, WE 2000aA rotated ellipsoidal angle density function improves estimation of foliage inclination distributions in forest canopiesAgric For Meteorol1001924CrossRefGoogle Scholar
  65. 65.
    Thomas, SC, Winner, WE 2000bLeaf area index of an old-growth Douglas-fir forest estimated from direct structural measurements in the canopyCan J For Res30192230CrossRefGoogle Scholar
  66. 66.
    Van Pelt, R, North, MP 1996Analyzing canopy structure in Pacific Northwest old-growth forests with a stand-scale crown modelNorthwest Sci70(special issue1531Google Scholar
  67. 67.
    Van Pelt, R, Franklin, JF 2000Influences of canopy structure on the understory environment in tall old-growth forestsCan J For Res30123145CrossRefGoogle Scholar
  68. 68.
    Weiss, SB 2000Spatial and temporal structure of insolation in an old-growth coniferous forestCan J For Res30195364CrossRefGoogle Scholar
  69. 69.
    Williams M, Rastetter EB, Fernandes DN, Goulden ML, Wofsy SC, Shaver GR, Melillo JM, and others. 1996. Modelling the soil–plant–atmosphere continuum in a Quercus-Acer stand at Harvard Forest: the regulation of stomatal conductance by light, nitrogen and soil/plant hydrologic properties. Plant Cell Environ 19:911–27.Google Scholar
  70. 70.
    Winner WE Thomas SC, Berry J, Bond BJ, Cooper CE, Hinckley TM, Ehleringer JR, and others. 2004. Canopy carbon gain and water use: analysis of old-growth conifers in the Pacific Northwest. Ecosystems 7:Google Scholar

Copyright information

© Springer-Verlag New York, Inc. 2004

Authors and Affiliations

  • Geoffrey G. Parker
    • 1
    Email author
  • Mark E. Harmon
    • 2
  • Michael A. Lefsky
    • 3
  • Jiquan Chen
    • 4
  • Robert Van Pelt
    • 5
  • Stuart B. Weis
    • 6
  • Sean C. Thomas
    • 7
  • William E. Winner
    • 8
  • David C. Shaw
    • 9
  • Jerry F. Frankling
    • 5
  1. 1.Smithsonian Environmental Research CenterEdgewaterUSA
  2. 2.Department of Forest ScienceOregon State UniversityCorvallisUSA
  3. 3.Department of Forest ScienceColorado State UniversityFort CollinsUSA
  4. 4.Landscape Ecology and Ecosystem ScienceUniversity of ToledoToledoUSA
  5. 5.College of Forest ResourcesUniversity of WashingtonSeattleUSA
  6. 6.27 Bishop LaneMenlo ParkUSA
  7. 7.Faculty of ForestryUniversity of TorontoTorontoCanada
  8. 8.Department of Botany and Plant PathologyOregon State UniversityCorvallisUSA
  9. 9.Wind River Canopy Crane Research FacilityCarsonUSA

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