, Volume 165, Issue 2, pp 533–544 | Cite as

Will the CO2 fertilization effect in forests be offset by reduced tree longevity?

  • Harald BugmannEmail author
  • Christof Bigler
Global change ecology - Original Paper


Experimental studies suggest that tree growth is stimulated in a greenhouse atmosphere, leading to faster carbon accumulation (i.e., a higher rate of gap filling). However, higher growth may be coupled with reduced longevity, thus leading to faster carbon release (i.e., a higher rate of gap creation). The net effect of these two counteracting processes is not known. We quantify this net effect on aboveground carbon stocks using a novel combination of data sets and modeling. Data on maximum growth rate and maximum longevity of 141 temperate tree species are used to derive a relationship between growth stimulation and changes in longevity. We employ this relationship to modify the respective parameter values of tree species in a forest succession model and study aboveground biomass in a factorial design of growth stimulation × reduced maximum longevity at multiple sites along a climate gradient from the cold to the dry treeline. The results show that (1) any growth stimulation at the tree level leads to a disproportionately small increase of stand biomass due to negative feedback effects, even in the absence of reduced longevity; (2) a reduction of tree longevity tends to offset the growth-related biomass increase; at the most likely value of reduced longevity, the net effect is very close to zero in most multi- and single-species simulations; and (3) when averaging the response across all sites to mimic a “landscape-level” response, the net effect is close to zero. Thus, it is important to consider ecophysiological responses with their linkage to demographic processes in forest trees if one wishes to avoid erroneous inference at the ecosystem level. We conclude that any CO2 fertilization effect is quite likely to be offset by an associated reduction in the longevity of forest trees, thus strongly reducing the carbon mitigation potential of temperate forests.


Carbon storage Climate change mitigation Growth stimulation Succession model ForClim 


  1. Arendt JD (1997) Adaptive intrinsic growth rates: an integration across taxa. Q Rev Biol 72:149–177CrossRefGoogle Scholar
  2. Backman G (1943) Wachstum und organische Zeit. Verlag Johann Ambrosius Barth, LepizigGoogle Scholar
  3. Bigler C, Bugmann H (2003) Growth-dependent tree mortality models based on tree rings. Can J For Res 33:210–221CrossRefGoogle Scholar
  4. Bigler C, Veblen TT (2009) Increased early growth rates decrease longevities of conifers in subalpine forests. Oikos 118:1130–1138CrossRefGoogle Scholar
  5. Bormann DB, Likens GE (1979) Pattern and process in a forested ecosystem. Springer, New YorkGoogle Scholar
  6. Botkin DB, Janak JF, Wallis JR (1972) Some ecological consequences of a computer model of forest growth. J Ecol 60:849–872CrossRefGoogle Scholar
  7. Bugmann H (1994) On the ecology of mountainous forests in a changing climate: a simulation study (PhD thesis, no. 10,638). Swiss Federal Institute of Technology, ZurichGoogle Scholar
  8. Bugmann H (1996) A simplified forest model to study species composition along climate gradients. Ecology 77:2055–2074CrossRefGoogle Scholar
  9. Bugmann H (1997) An efficient method for estimating the steady-state species composition of forest gap models. Can J For Res 27:551–556CrossRefGoogle Scholar
  10. Bugmann H (2001a) A review of forest gap models. Clim Change 51:259–305CrossRefGoogle Scholar
  11. Bugmann H (2001b) A comparative analysis of forest dynamics in the Swiss Alps and the Colorado Front Range. For Ecol Manag 145:43–55CrossRefGoogle Scholar
  12. Bugmann H (2003) Predicting the ecosystem effects of climate change. In: Canham CD, Lauenroth WK, Cole JS (eds) Models in ecosystem science. Princeton University Press, Princeton, pp 385–409Google Scholar
  13. Bugmann H, Cramer W (1998) Improving the behaviour of forest gap models along drought gradients. For Ecol Manag 103:247–263CrossRefGoogle Scholar
  14. Bugmann H, Solomon AM (1995) The use of a European forest model in North America: a study of ecosystem response to climate gradients. J Biogeogr 22:477–484CrossRefGoogle Scholar
  15. Bugmann H, Solomon AM (2000) Explaining forest composition and biomass across multiple biogeographical regions. Ecol Appl 10:95–114CrossRefGoogle Scholar
  16. Cramer W et al (2001) Global response of terrestrial ecosystem structure and function to CO2 and climate change: results from six dynamic global vegetation models. Glob Change Biol. 7:357–373Google Scholar
  17. Friedlingstein P et al (2006) Climate–carbon cycle feedback analysis: results from the C4MIP model intercomparison. J Clim 19:3337–3353Google Scholar
  18. Friend AD, Stevens AK, Knox RG, Cannell MGR (1997) A process-based, terrestrial biosphere model of ecosystem dynamics (Hybrid v3.0). Ecological Modelling 95:249–287CrossRefGoogle Scholar
  19. Gayler S, Grams TEE, Heller W, Treutter D, Priesack E (2008) A dynamical model of environmental effects on allocation to carbon-based secondary compounds in juvenile trees. Ann Bot 101:1089–1098CrossRefPubMedGoogle Scholar
  20. Granados J, Körner C (2002) In deep shade, elevated CO2 increases the vigor of tropical climbing plants. Global Change Biol 8:1109–1117Google Scholar
  21. Handa IT, Körner C, Hättenschwiler S (2005) A test of the treeline carbon limitation hypothesis by in situ CO2 enrichment and defoliation. Ecology 86:1288–1300CrossRefGoogle Scholar
  22. Harcombe PA (1987) Tree life tables. Bioscience 37:557–568CrossRefGoogle Scholar
  23. Jenkins MA, Pallardy SG (1995) The influence of drought on red oak group species growth and mortality in the Missouri Ozarks. Can J For Res 25:1119–1127Google Scholar
  24. Kimball BA, Pinter PJ, Garcia RL, LaMorte RL, Wall GW, Hunsaker DJ, Wechsung G, Wechsung F, Kartschall Th (1995) Productivity and water use of wheat under free-air CO2 enrichment. Glob Change Biol 1:429–442CrossRefGoogle Scholar
  25. Kimball BA, Idso SB, Johnson S, Rillig MC (2007) Seventeen years of carbon dioxide enrichment of sour orange trees: final results. Glob Change Biol 13:2171–2183CrossRefGoogle Scholar
  26. Kimmins JP (2004) Forest ecology. Prentice-Hall, Upper Saddle RiverGoogle Scholar
  27. Körner C (2004) Through enhanced tree dynamics carbon dioxide enrichment may cause tropical forests to lose carbon. Phil Trans R Soc Lond B 359:493–498CrossRefGoogle Scholar
  28. Körner C (2006) Plant CO2 responses: an issue of definition, time and resource supply. New Phytol 172:393–411CrossRefPubMedGoogle Scholar
  29. Körner C (2009) Responses of humid tropical trees to rising CO2. Annu Rev Ecol Evol Syst 40:61–79CrossRefGoogle Scholar
  30. Körner C, Asshoff R, Bignucolo O, Hättenschwiler S, Keel SG, Pelaez-Riedl S, Pepin S, Siegwolf RTW, Zotz G (2005) Carbon flux and growth in mature deciduous forest trees exposed to elevated CO2. Science 309:1360–1362CrossRefPubMedGoogle Scholar
  31. Körner C, Morgan JA, Norby R (2007) CO2 fertilisation: when, where, how much? In: Canadell JG, Pataki D, Pitelka LF (eds) Terrestrial ecosystems in a changing world. Springer, Berlin, pp 9–21Google Scholar
  32. Ladeau SL, Clark JS (2006) Elevated CO2 and tree fecundity: the role of tree size, interannual variability, and population heterogeneity. Glob Change Biol 12:822–833CrossRefGoogle Scholar
  33. Laurance WF, Nascimento HEM, Laurance SG, Condit R, D’Angelo S, Andrade A (2004) Inferred longevity of Amazonian rainforest trees based on a long-term demographic study. For Ecol Manag 190:131–143CrossRefGoogle Scholar
  34. Lavola A, Julkunen-Tiitto R (1994) The effect of elevated carbon dioxide and fertilization on primary and secondary metabolites in birch Betula pendula (Roth). Oecologia 99:315–321CrossRefGoogle Scholar
  35. Lexer MJ, Hönninger K (2001) A modified 3D-patch model for spatially explicit simulation of vegetation composition in heterogeneous landscapes. For Ecol Manag 144:43–65CrossRefGoogle Scholar
  36. Litvak ME, Constable JVH, Monson RK (2002) Supply and demand processes as controls over needle monoterpene synthesis and concentration in Douglas fir [Pseudotsuga menziesii (Mirb.) Franco]. Oecologia 132:382–391CrossRefGoogle Scholar
  37. Loehle C (1988) Tree life history strategies: the role of defenses. Can J For Res 18:209–222Google Scholar
  38. Luo ZB, Calfapietra C, Scarascia-Mugnozza G, Liberloo M, Polle A (2008) Carbon-based secondary metabolites and internal nitrogen pools in Populus nigra under free air CO2 enrichment (FACE) and nitrogen fertilization. Plant Soil 304:45–57Google Scholar
  39. Malhi Y, Meir P, Brown S (2002) Forests, carbon and global climate. Phil Trans R Soc Lond B 360:1567–1591CrossRefGoogle Scholar
  40. Moore AD (1989) On the maximum growth equation used in forest gap simulation models. Ecological Modelling 45:63–67CrossRefGoogle Scholar
  41. Moore DJP, Aref S, Ho RM, Pippen JS, Hamilton JG, DeLucia EH (2006) Annual basal area increment and growth duration of Pinus taeda in response to eight years of free-air carbon dioxide enrichment. Glob Change Biol 12:1367–1377CrossRefGoogle Scholar
  42. Norby RJ, Wullschleger SD, Gunderson CA, Johnson DW, Ceulemans R (1999) Tree responses to rising CO2 in field experiments: implications for the future forest. Plant Cell Environ 22:683–714CrossRefGoogle Scholar
  43. Norby RJ, Ledford J, Reilly CD, Miller NE, O’Neill EG (2004) Fine-root production dominates response of a deciduous forest to atmospheric CO2 enrichment. Proc Natl Acad Sci USA 101:9689–9693CrossRefPubMedGoogle Scholar
  44. Norby RJ et al (2005) Forest response to elevated CO2 is conserved across a broad range of productivity. Proc Natl Acad Sci USA 102:18052–18056Google Scholar
  45. Oren R, Ellsworth DS, Johnsen KH, Phillips N, Ewers BE, Maier C, Schäfer KVR, McCarthy H, Hendrey G, McNulty SG, Katul GG (2001) Soil fertility limits carbon sequestration by forest ecosystems in a CO2-enriched atmosphere. Nature 411:469–472CrossRefPubMedGoogle Scholar
  46. Philips OL, Gentry AH (1994) Increasing turnover through time in tropical forests. Science 263:954–957CrossRefGoogle Scholar
  47. Philips OL et al (2004) Pattern and process in Amazon tree turnover, 1976–2001. Phil Trans R Soc Lond B 359:381–407Google Scholar
  48. Prentice IC, Cramer W, Harrison SP, Hickler T, Lucht W, Sitch S, Smith B, Sykes MT (2007) Dynamic global vegetation modelling: quantifying terrestrial ecosystem responses to large-scale environmental change. In: Canadell JG, Pataki D, Pitelka LF (eds) Terrestrial ecosystems in a changing world. Springer, Berlin, pp 175–192Google Scholar
  49. Purves DW, Lichstein JW, Pacala SW (2007) Crown plasticity and competition for canopy space: a new spatially implicit model parameterized for 250 North American tree species. PLoS ONE 2(9):e870CrossRefPubMedGoogle Scholar
  50. R Development Core Team (2008) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna.
  51. Reynolds JF, Bugmann H, Pitelka LF (2001) How much physiology is needed in forest gap models for simulating long-term vegetation response to global change? Challenges, limitations, and potentials. Clim Change 51:541–557CrossRefGoogle Scholar
  52. Schäfer KVR, Oren R, Ellsworth DS, Lai CT, Herrick JD, Finzi AC, Richter DD, Katul GG (2003) Exposure to an enriched CO2 atmosphere alters carbon assimilation and allocation in a pine forest ecosystem. Glob Change Biol 9:1378–1400CrossRefGoogle Scholar
  53. Schulman E (1954) Longevity under adversity in conifers. Science 119:396–399CrossRefPubMedGoogle Scholar
  54. Shao G, Bugmann H, Yan X (2001) A comparative analysis of the structure and behavior of three forest gap models at sites in northeastern China. Clim Change 51:389–413CrossRefGoogle Scholar
  55. Shugart HH (1984) A theory of forest dynamics: the ecological implications of forest succession models. Springer, New YorkGoogle Scholar
  56. Shugart HH (1998) Terrestrial ecosystems in a changing environment. Cambridge University Press, CambridgeGoogle Scholar
  57. Shugart HH, Emanuel WR (1985) Carbon dioxide increase: the implications at the ecosystem level. Plant Cell Environ 8:381–386CrossRefGoogle Scholar
  58. Vieira S, Trumbore S, Camargo PB, Selhorst D, Chambers JQ, Higuchi N, Martinelli LA (2005) Slow growth rates of Amazonian trees: consequences for carbon cycling. Proc Natl Acad Sci USA 102:18502–18507CrossRefPubMedGoogle Scholar
  59. Waring RH (1987) Characteristics of trees predisposed to die. Bioscience 37:569–573CrossRefGoogle Scholar
  60. Wright SJ, Calderon O, Hernandez A, Paton S (2004) Are lianas increasing in importance in tropical forests? A 17-year record from Panama. Ecology 85:484–489CrossRefGoogle Scholar
  61. Wunder J, Brzeziecki B, Żybura H, Reineking B, Bigler C, Bugmann H (2008) Growth–mortality relationships as indicators of life-history strategies: a comparison of nine tree species in unmanaged European forests. Oikos 117:815–828CrossRefGoogle Scholar
  62. Wyckoff PH, Clark JS (2002) The relationship between growth and mortality for seven co-occurring tree species in the southern Appalachian Mountains. J Ecol 90:604–615CrossRefGoogle Scholar
  63. Zotz G, Cueni N, Körner C (2006) In situ growth stimulation of a temperate zone liana (Hedera helix) in elevated CO2. Funct Ecol 20:763–769CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2010

Authors and Affiliations

  1. 1.Laboratory of Tree-Ring ResearchUniversity of ArizonaTucsonUSA
  2. 2.Forest Ecology, Department of Environmental Sciences, Institute of Terrestrial EcosystemsETH ZurichZurichSwitzerland

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