, Volume 11, Issue 3, pp 426–441 | Cite as

Gaps and Soil C Dynamics in Old Growth Northern Hardwood–Hemlock Forests



Old growth forest soils are large C reservoirs, but the impacts of tree-fall gaps on soil C in these forests are not well understood. The effects of forest gaps on soil C dynamics in old growth northern hardwood–hemlock forests in the upper Great Lakes region, USA, were assessed from measurements of litter and soil C stocks, surface C efflux, and soil microbial indices over two consecutive growing seasons. Forest floor C was significantly less in gaps (19.0 Mg C ha−1) compared to gap-edges (39.5 Mg C ha−1) and the closed forest (38.0 Mg C ha−1). Labile soil C (coarse particulate organic matter, cPOM) was significantly less in gaps and edges (11.1 and 11.2 Mg C ha−1) compared to forest plots (15.3 Mg C ha−1). In situ surface C efflux was significantly greater in gaps (12.0 Mg C ha−1 y−1) compared to edges and the closed forest (9.2 and 8.9 Mg C ha−1 y−1). Microbial biomass N (MBN) was significantly greater in edges (0.14 Mg N ha−1) than in the contiguous forest (0.09 Mg N ha−1). The metabolic quotient (qCO2) was significantly greater in the forest (0.0031 mg CO2 h−1 g−1/mg MBC g−1) relative to gaps or edges (0.0014 mg CO2 h−1 g−1/mg MBC g−1). A case is made for gaps as alleviators of old growth forest soil C saturation. Relative to the undisturbed closed forest, gaps have significantly less labile C, significantly greater in situ surface C efflux, and significantly lower decreased qCO2 values.


forest gap microbial biomass microbial metabolic quotient (qCO2microbial biomass to organic C ratio old growth forest particulate organic matter surface C efflux 


  1. Allen SE. 1989. Chemical Analysis of Ecological Materials. 2nd edition. Blackwell Scientific Publications, Oxford, UK.Google Scholar
  2. Alvarez CR, Alvarez R, Grigera MS, Lavado RS. 1998. Associations between organic matter fractions and the active soil microbial biomass. Soil Biol Biochem 30:767–773.CrossRefGoogle Scholar
  3. Anderson JPE, Domsch KH. 1978. A physiological method for the quantitative measurement of microbial biomass in soils. Soil Biol Biochem 10:215–221.CrossRefGoogle Scholar
  4. Anderson TH, Domsch KH. 1985. Maintenance carbon requirements of actively-metabolizing microbial populations under in situ conditions. Soil Biol Biochem 17:197–203.CrossRefGoogle Scholar
  5. Anderson TH, Domsch KH. 1989. Ratios of microbial biomass carbon to total organic carbon in arable soils. Soil Biol Biochem 21:471–479.CrossRefGoogle Scholar
  6. Bauhus J, Bartsch N. 1995. Mechanisms for carbon and nutrient release and retention in beech forest gaps. Plant Soil 168–169:579–584.CrossRefGoogle Scholar
  7. Bauhus J. 1996. C and N mineralization in an acid forest soil along a gap-stand gradient. Soil Biol Biochem 28:923–932.CrossRefGoogle Scholar
  8. Bhatti JS, Apps MJ, Tarnocai C. 2002. Estimates of soil organic carbon stocks in central Canada using three different approaches. Can J For Res 32:805–812.CrossRefGoogle Scholar
  9. Bockheim JG. 2003. Genesis of bisequal soils on acidic drift in the Upper Great Lakes region, U.S.A. Soil Sci Soc Am J 67:612–619.CrossRefGoogle Scholar
  10. Brady NC, Weil RR. 2002. The nature and properties of soils. 13th Ed. Prentice Hall, New Jersey, USA.Google Scholar
  11. Bremer E, Janzen HH, Johnston AM. 1994. Sensitivity of total, light, and mineralizable organic matter to management practices in a Lethbridge soil. Can J Soil Sci 74:131–138.Google Scholar
  12. Brookes PC, Landman A, Pruden G, Jenkinson DS. 1985. Chloroform fumigation and the release of soil nitrogen: A rapid direct extraction method to measure microbial biomass nitrogen in soil. Soil Biol Biochem 17:837–842.CrossRefGoogle Scholar
  13. Brown S, Lugo AE. 1982. The storage and production of organic matter in tropical forests and their role in the global carbon cycle. Biotropica 14:161–187.CrossRefGoogle Scholar
  14. Brumme R. 1995. Mechanisms of carbon and nutrient release and retention in beech forest gaps. Plant Soil 168–169:593–600.CrossRefGoogle Scholar
  15. Buchmann N, Schulze E. 1999. Net CO2 and H2O fluxes of terrestrial ecosystems. Global Biogeochem Cycles 13:751–760.CrossRefGoogle Scholar
  16. Bunnell FL, Tait DEN, Flanagan PW, Van Cleve K. 1977. Microbial respiration and substrate weight loss-I: A general model of the influences of abiotic variables. Soil Biol Biochem 9:33–40.CrossRefGoogle Scholar
  17. Buyanovsky GA, Aslam M, Wagner GH. 1994. Carbon turnover in soil physical fractions. Soil Sci Soc Am J 58:1167–1173.CrossRefGoogle Scholar
  18. Cabrera ML, Beare MH. 1993. Alkaline persulfate oxidation for determining total nitrogen in microbial biomass estimates. Soil Sci Soc Am J 57:1007–1012.CrossRefGoogle Scholar
  19. Cambardella CA, Elloitt ET. 1992. Particulate soil organic matter changes across a grassland cultivation sequence. Soil Sci Soc Am J 56:777–783.CrossRefGoogle Scholar
  20. Cambardella CA, Elloitt ET. 1993. Carbon and nitrogen distribution in aggregates from cultivated and native grassland soils. Soil Sci Soc Am J 57:1071–1076.CrossRefGoogle Scholar
  21. Campbell CA, Bowren KE, Schnitzer M, Zentner RP, Townley-Smith L. 1991. Effect of crop rotations and fertilization on soil biochemical properties in a thick Black Chernozem. Can J Soil Sci 71: 377–387.Google Scholar
  22. Canham CD, Loucks OL. 1984. Catastrophic wind-throw in the presettlement forests of Wisconsin. Ecology 65:803–809.CrossRefGoogle Scholar
  23. Catroux G, Schnitzer M. 1987. Chemical, spectroscopic and biological characteristics of the organic matter in particle size fractions separated from an Aquoll. Soil Sci Soc Am J 51:1200–1207.CrossRefGoogle Scholar
  24. Cerovic ZG, Godulas Y, Gorbunov M, Briantais J-M, Camenen L, Moya I, Winkler JP, Cherry RS, Schlesinger WH. 1996. The Q10 relationship of microbial respiration in a temperate forest soil. Soil Biol Biochem 28:1067–1072.CrossRefGoogle Scholar
  25. Coleman DC, Crossley Jr DA, Hendrix PF. 2004. Fundamentals of soil ecology. 2nd Edition. Elsevier Academic Press, San Diego, CA, USA.Google Scholar
  26. Collins BS, Pickett STA. 1987. Response of herb layer cover to experimental canopy gaps. American Midlander Naturalist 119:282–290.CrossRefGoogle Scholar
  27. Curtin D, Wen G. 1999. Organic matter fractions contributing to soil nitrogen mineralization potential. Soil Sci Soc Am J 63:1361–1365.CrossRefGoogle Scholar
  28. Curtis JT. 1959. The vegetation of Wisconsin. University of Wisconsin Press, Madison.Google Scholar
  29. Dahir SE, Lorimer CG. 1996. Variation in canopy gap formation among developmental stages of northern hardwood stands. Can J For Res 26:1875–1892.CrossRefGoogle Scholar
  30. Dalal RC, Mayer RJ. 1986. Long term trends in fertility of soils under continuous cultivation and cereal cropping in southern Queensland. III. distribution and kinetics of soil organic carbon in particle-size fractions. Austr J Soil Res 24:301–309.CrossRefGoogle Scholar
  31. Davis MR, Calcote RR, Sugita S, Takahura H. 1998. Patch invasion and origin of a hemlock-hardwoods forest mosaic. Ecology 79:2641–2659.Google Scholar
  32. Denslow JS, Ellison AM, Sanford RE. 1998. Treefall gap size effects on above- and below-ground processes in a tropical wet forest. Ecology 86:597–609.CrossRefGoogle Scholar
  33. Drinkwater LE, Cambardella CA, Reeder J, Rice CW. 1996. Potentially mineralizable nitrogen as an indicator of biologically active soil nitrogen. In: Doran JW, Jones AJ (Eds) Methods for Assessing Soil Quality. Soil Science Society of America, Inc., Madison, Wisconsin pp. 217–229.Google Scholar
  34. Edwards NT. 1982. The use of soda lime for measuring respiration rates in terrestrial systems. Pedobiologia 23:321–330.Google Scholar
  35. Elliot ET, Cambardella CA. 1991. Physical separation of soil organic matter. Agric Ecosyst Environ 34:407–419.CrossRefGoogle Scholar
  36. Field CB, Kaduk J. 2004. The carbon balance of an old-growth forest: building across approaches. Ecosystems 7:525–533.CrossRefGoogle Scholar
  37. Fredeen AL, Bois CH, Janzen DT, Sanborn PT. 2005. Comparison of coniferous forest carbon stocks between old-growth and young second-growth forests on two soil types in central British Columbia, Canada. Can J For Res 35:1411–1421.CrossRefGoogle Scholar
  38. Frelich LE, Calcote RR, Davis MB. 1993. Patch formation and maintenance in an old-growth hemlock hardwood forest. Ecology 74:513–527.CrossRefGoogle Scholar
  39. Frelich LE, Lorimer CG. 1991. Natural disturbance regimes in hemlock-hardwood forests of the upper Great Lakes region. Ecological Monographs 61:145–161.CrossRefGoogle Scholar
  40. Gardner WH. 1965. Water content. In: Black CA, others, Eds. Methods of soil analysis. Part II. Agron. Monogr. 9. Madison, WI, USA: ASA.Google Scholar
  41. Goodburn JM, Lorimer CG. 1999. Population structure in old-growth and managed northern hardwoods: an examination of the balanced diameter distribution concept. For Ecol Man 118:11–29.CrossRefGoogle Scholar
  42. Gregorich EG, Ellert BH. 1993. Light fraction and macroorganic matter in mineral soils. In: Carter MR (Ed) Soil sampling and methods of analysis. Lewis Publishers, CRC Press Inc., Boca Raton, FL pp. 397–407.Google Scholar
  43. Grogan P. 1998. CO2 flux measurement using soda lime: correction for water formed during CO2 adsorption. Ecology 79:1467–1468.CrossRefGoogle Scholar
  44. Halvorston JJ, Smith JL, Franz EH. 1991. Lupine influences on soil carbon, nitrogen, and microbial activity in developing ecosystems at Mount St. Helens. Oecologia 87:162–170.CrossRefGoogle Scholar
  45. Harmon ME, Bible K, Ryan MG, Shaw DG, Chen H, Klopatek J, Li X. 2004. Production, respiration, and overall carbon balance in an old-growth Pseudotsuga-Tsuga forest ecosystem. Ecosystems 7:498–512.Google Scholar
  46. Hart SC, Firestone MK. 1991. Forest floor-mineral soil interactions in the internal nitrogen cycle of an old-growth forest. Biogeochemistry 12:103–127.CrossRefGoogle Scholar
  47. Hart SC, Nason GE, Myrold DD, Perry DA. 1994. Dynamics of gross nitrogen transformation in an old-growth forest: the carbon connection. Ecology 75:880–891.CrossRefGoogle Scholar
  48. Holmes WE, Zak DR. 1999. Soil microbial control of nitrogen loss following clear-cut harvest in northern hardwood ecosystems. Ecol Appl 9:202–215.CrossRefGoogle Scholar
  49. Houghton RA, Hackler JL, Lawerence KT. 1999. The US carbon budget: contributions from land-use change. Science 285:574–578.PubMedCrossRefGoogle Scholar
  50. Insam H, Domsch KH. 1988. Relationship between soil organic carbon and microbial biomass on chronosequences of reclamation sites. Microb Ecol 15:177–188.CrossRefGoogle Scholar
  51. Insam H, Hasselwandter K. 1989. Metabolic quotient of the soil microflora in relation to plant succession. Oecologia 79:174–178.CrossRefGoogle Scholar
  52. Insam H, Parkinson D, Domsch KH. 1989. Influence of macroclimate on soil microbial biomass. Soil Biol Biochem 21:211–221.CrossRefGoogle Scholar
  53. Jenkinson DS, Rayner JH. 1977. The turnover of soil organic matter in some of the Rothamsted classical experiments. Soil Sci 123:298–305.CrossRefGoogle Scholar
  54. Joergensen RG, Mueller T. 1996. The fumigation-extraction method to estimate soil microbial biomass: calibration of the kEN value. Soil Biol Biochem 28:33–37.CrossRefGoogle Scholar
  55. Kaiser EA, Mueller T, Joergensen RG, Insam H, Heinemeyer O. 1992. Evaluation of methods to estimate the soil microbial biomass and the relationship with soil texture and organic matter. Soil Biol Biochem 27:675–683.CrossRefGoogle Scholar
  56. Keith H, Wong SC. 2006. Measurement of soil CO2 efflux using soda lime absorption: both quantitative and reliable. Soil Biol Biochem 38: 1121–1131.CrossRefGoogle Scholar
  57. LECO CNS. 2000. Carbon analyzer. University of Wisconsin Soil and Plant Analysis Laboratory.Google Scholar
  58. McClure JW, Lee TD, Leak WB. 2000. Gap capture in northern hardwoods: patterns of establishment an height growth in four species. Forest Ecol Man 127:181–189.CrossRefGoogle Scholar
  59. McGill WB, Cannon K, Robertson JA, Cook FD. 1986. Dynamics of soil microbial biomass and water soluble organic C in Breton L after 50 years of cropping to two rotations. Can J Soil Sci 66:1–19.CrossRefGoogle Scholar
  60. Mladenoff DJ. 1987. Dynamics of nitrogen mineralization and nitrification in hemlock and hardwood treefall gaps. Ecology 68:1171–1180.CrossRefGoogle Scholar
  61. Monteith JL, Steicz G, Yabuki K. 1964. Crop photosynthesis and the flux of carbon dioxide below the canopy. J Appl Ecol 1:321–337.CrossRefGoogle Scholar
  62. National Oceanic, Atmospheric Administration (NOAA). 2007. Climatological data national summary. Environmental Data Service National Climatic Center, North Carolina.Google Scholar
  63. Nohrstedt HO, Arnebrant K, Baath E, Soderstrom B. 1989. Changes in carbon content, respiration rate, ATP content and microbial biomass in nitrogen-fertilised pine forest soils in central Sweden. Can J For Res 19:323–328.CrossRefGoogle Scholar
  64. Odum EP. 1969. The strategy of ecosystem development. Science 164: 262–270.PubMedCrossRefGoogle Scholar
  65. Odum EP. 1985. Trends in stressed ecosystems. Bioscience 35: 419–422.CrossRefGoogle Scholar
  66. Onset Computer Corporation. 2003. Pocasset. Massachutes, USA.Google Scholar
  67. Parkin TB, Doran JW, Franco-Vizcaino E. 1996. Field and laboratory tests of soil respiration. In: Doran JW, Jones AJ (Eds) Methods for Assessing Soil Quality. Soil Science Society of America, Inc., Madison, Wisconsin pp. 231–245.Google Scholar
  68. Parsons WFJ, Knight DH, Miller SL. 1994a. Root gap dynamics and forest disturbance theory in lodgepole pine forests: nitrogen transformations in gaps of different size. Ecol Appl 4:354–362.CrossRefGoogle Scholar
  69. Parsons WFJ, Knight DH, Miller SL. 1994b. Root gap dynamics in lodgepole pine forests: ectomycorrhizal and nonmycorrhizal fine root activity after experimental gap formation. Can J For Res 24:1531–1538.CrossRefGoogle Scholar
  70. Parton WJ, Schimel DS, Cole CV, Ojima DS. 1988. Analysis of factors controlling soil organic matter levels in great plains grasslands. Soil Sci. Soc. Am. J 51:1173–1179.CrossRefGoogle Scholar
  71. Pastor J, Broschart M. 1990. The spatial pattern of a northern conifer hardwood landscape. Landscape Ecology 4:55–68.CrossRefGoogle Scholar
  72. Paul EA, Juma NG. 1981. Mineralization and immobilization of soil nitrogen by microorganisms. In: Clark FE, Rosswall T, Eds. Terrestrial nitrogen cycles. Ecological Bulletins 33, Stockholm. pp 179–95.Google Scholar
  73. Paustian K, Collins HP, Paul EA. 1997. Management controls on soil carbon. In: Paul EA, Paustian K, Elliot ET, Cole CV (Eds) Soil organic matter in temperate agroecosystems. CRC Press, Boca Raton, FL, USA pp. 15–49.Google Scholar
  74. Paw U KT, Falk M, Suchanek TH, Ustin SL, Chen J, Park Y, Winner WW, Thomas SC, Hsiao TC, Shaw RH, King TS, Pyles RD, Schroeder M, Matista AA. 2004. Carbon dioxide exchange between an old-growth forest and the atmosphere. Ecosystems 7:513–524.Google Scholar
  75. Phillips DW, McCulloch JAW. 1972. The climate of the Great Lakes Basin. Climatological Studies Number 20 Atmospheric Environment Service Environment Canada, Toronto.Google Scholar
  76. Poulson TL, Platt WJ. 1996. Replacement patterns of beech and sugar maple in Warren Woods, Michigan. Ecology 77:1234–1253.CrossRefGoogle Scholar
  77. Prevost M. 2004. Predicting soil properties from organic matter content following mechanical site preparation of forest soils. Soil Sci Soc Am J 68: 943–949.CrossRefGoogle Scholar
  78. Quiroga AR, Buschiazzo DE, Peinemann N. 1996. Soil organic matter particle size fractions in soils of the semiarid Argentinean pampas. Soil Sci 161:104–108.CrossRefGoogle Scholar
  79. Ross DJ. 1992. Influence of sieve mesh size on estimates of microbial carbon and nitrogen by fumigation-extraction procedures in soils under pasture. Soil Biol Biochem 24:343–350.CrossRefGoogle Scholar
  80. Runkle JR. 1992. Guidelines and sample protocol for sampling forest gaps. General Techniques Reproduction PNW GTR 283 U.S. Department of Agriculture, Forest Service, Pacific Northwest Resource Station.Google Scholar
  81. SAS. 2005. SAS Institute Inc. Cary, NC, USA.Google Scholar
  82. Schaetzl RJ, Follmer LR. 1990. Longevity of treethrow microtopography: implications for mass wasting. Geomorphology 3:113–123.CrossRefGoogle Scholar
  83. Scharenbroch BC, Bockheim JG. 2007. Impacts of forest gaps on soil properties and processes in old growth northern hardwood–hemlock forests. Plant Soil 294:219–233.CrossRefGoogle Scholar
  84. Schulte EE, Kaufman C, Peter JB. 1991. The influence of sample size and heating time on soil weight loss-on-ignition. Commun Soil Sci Plant Anal 22:159–166.CrossRefGoogle Scholar
  85. Sierra J. 1996. Nitrogen mineralisation and its error of estimation under field conditions related to the light fraction of soil organic matter. Austr J Soil Res 34:755–767.CrossRefGoogle Scholar
  86. Sikora LJ, Stott DE. 1996. Soil organic carbon and nitrogen. In: Doran JW, Jones AJ (Eds) Methods for Assessing Soil Quality. Soil Science Society of America, Inc., Madision, Wisconsin pp. 157–167.Google Scholar
  87. Singh JS, Raghubanshi AS, Singh RS, Srivastasta SL. 1989. Microbial biomass acts as a source of plant nutrients in dry tropical forests and savanna. Nature 338:449–500.CrossRefGoogle Scholar
  88. Six J, Contant T, Paul EA, Paustian K. 2002. Stabilization mechanisms of soil organic matter: Implications for C-saturation of soils. Plant Soil 241:155–176.CrossRefGoogle Scholar
  89. Smithwick EAH, Harmon ME, Remillard SM, Acker SA, Franklin JF. 2002. Potential upper bounds of carbon stores in forests of the Pacific Northwest. Ecol Appl 12:1303–1317.CrossRefGoogle Scholar
  90. Solberg ED, Nyborg M, Izaurralde RC, Malhi SS, Janzen HH, Molina-Ayala M. 1997. Carbon storage in soils under continuous cereal grain cropping: N fertilizer and straw. In: Lal R, Kimble JM, Follett RF, Stewart BA (Eds) Management of carbon sequestration in soil. CRC Press, Boca Raton, FL, USA pp. 235–254.Google Scholar
  91. Solomon D, Lehmann J, Zech W. 2000. Land use effects on soil organic matter properties of chronic Luvisols in semi-arid northern Tanzania: carbon, nitrogen, lignin and carbohydrates. Agr Ecosys Environ 78:203–213.CrossRefGoogle Scholar
  92. Sparling GP, Feltman CW, Reynolds J, West AW, Singleton P. 1990. Estimation of soil microbial C by a fumigation-extraction method: use on soils of high organic matter content, and a reassessment of the kEC factor. Soil Biol Biochem 22:301–307.CrossRefGoogle Scholar
  93. Suchanek TH, Mooney HA, Franklin JF, Gucinski H, Ustin SL. 2004. Carbon dynamics of an old-growth forest. Ecosystems 7:421–426.CrossRefGoogle Scholar
  94. Tissen H, Stewart JWB. 1993. Particle-size fractions and their use in studies of soil organic matter composition in size fractions. Soil Sci Soc Am J 47:509–514.CrossRefGoogle Scholar
  95. Turchenek LW, Oades JM. 1979. Fractionation of organomineral complexes by sedimentation and density techniques. Geoderma 21:311–343.CrossRefGoogle Scholar
  96. Tyrrell LE, Crow TR. 1994. Structural characteristics of old-growth hemlock-hardwood forests in relation to age. Ecology 75:370–386.CrossRefGoogle Scholar
  97. USDA NRCS. 2006. Keys to soil taxonomy, 10th edn. Madison, WI: United States Department of Agriculture, National Resources Conservation Service. National Cooperative Soil Survey.Google Scholar
  98. Wardle DA, Ghani A. 1995. A critique of the microbial metabolic quotient (qCO2) as a bioindicator of disturbance and ecosystem development. Soil Biol Biochem 27:1601–1610.CrossRefGoogle Scholar
  99. Wardle DA. 1993. Changes in the microbial biomass and metabolic quotient during leaf litter succession in some New Zealand forest and scrubland ecosystems. Functional Ecol 7:346–355.CrossRefGoogle Scholar
  100. Wolters V. 1991. Biological processes in two beech forest soils treated with simulated acid rain—a laboratory experiment with Isotoma tigrina(Insecta, Collembola). Soil Biol Biochem 23:381–390.CrossRefGoogle Scholar
  101. Woods KD. 2000. Long-term change and spatial pattern in a late-successional hemlock-northern hardwood forest. Ecology 88:267–282.CrossRefGoogle Scholar
  102. Woods KD. 2004. Intermediate disturbance in a late-successional hemlock-northern hardwood forest. Ecology 92:464–476.CrossRefGoogle Scholar
  103. Zhang Q, Zak JC. 1995. Effects of gap size on litter decomposition and microbial activity in a subtropical forest. Ecology 76:2196–2204.CrossRefGoogle Scholar
  104. Zhou G, Liu S, Li Z, Zhang D, Tang X, Zhou C, Yan J, Mo J. 2006. Old-growth forests can accumulate carbon in soils. Science 314:1417.PubMedCrossRefGoogle Scholar
  105. Zibilske LM. 1994. Carbon mineralization. In: Weaver RW, Angle JS, Bottomley PS (Eds) Methods of soil analysis: Part 2. Microbiological and biochemical properties. Soil Science Society of America, Madison, WI, USA pp. 835–864.Google Scholar

Copyright information

© Springer Science+Business Media, LLC 2008

Authors and Affiliations

  1. 1.The Morton ArboretumLisleUSA
  2. 2.Department of Soil ScienceUniversity of WisconsinMadisonUSA

Personalised recommendations