Plant Ecology

, Volume 187, Issue 2, pp 179–187 | Cite as

Contrasted Responses of Two Understorey Species to Direct and Indirect Effects of a Canopy Gap



Positive associations between adult trees and understorey species have been explained either by direct or indirect facilitation. We tested both models by comparing the performance of two understorey species with contrasted stress-tolerance abilities Galium odoratum and Deschampsia flexuosa. Individuals of both species were transplanted in the four combinations of two treatments (gap and removal of an herbaceous competitor, Molinia caerulea). Our experiment demonstrated that direct facilitation of adult trees may explain the restricted occurrence of the shade-demanding Galium within closed forest communities. In contrast, the shade-tolerant Deschampsia was subjected to additional competition within the forest, likely because adult trees had a higher negative effect on light availability and a similar negative effect on nitrogen availability within the forest than did Molinia in the gaps.


Competition intensity Forest communities Indirect effects Light Molinia caerulea Nitrogen 


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  1. M.P. Austin, Continuum conceptordination methods, and niche theory. Annu. Rev. Ecol. Syst. 16 (1985) 39-61CrossRefGoogle Scholar
  2. F.A. Bazzaz, The physiological ecology of plant succession. Annu. Rev. Ecol. Syst. 10 (1979) 351-371CrossRefGoogle Scholar
  3. E.L. Berlow, Strong effects of weak interactions in ecological communities. Nature 398 (1999) 330-334CrossRefGoogle Scholar
  4. F.H. Bormann and G.E. Likens, Pattern and Process in a Forested Ecosystem. New-York: Springer Verlag (1994).Google Scholar
  5. J. Braun-Blanquet, G.D. Fuller and H.S. Conard, Plant Sociology. The Study of Plant Communities. New-York: Mc-Graw-Hill edn (1932).Google Scholar
  6. R.W. Brooker and T.V. Callaghan, The balance between positive and negative plant interactions and its relationship to environmental gradients: a model. Oikos 81 (1998) 196-207Google Scholar
  7. J.F. Bruno, J.J. Stachowicz and M.D. Bertness, Inclusion of facilitation into ecological theory. Trends Ecol. Evol. 18 (2003) 119-125CrossRefGoogle Scholar
  8. R.M. Callaway, Positive interactions among plants. Bot. Rev. 61 (1995) 306-349CrossRefGoogle Scholar
  9. R.M. Callaway, Positive interactions in plant communities and the individualistic-continuum concept. Oecologia 112 (1997) 143-149CrossRefGoogle Scholar
  10. R.M. Callaway, E.H. Delucia, D. Moore, R. Nowak and W.H. Schlesinger, Competition and facilitation: contrasting effects of Artemisia tridentata on desert vs montane pines. Ecology 77 (1996) 2130-2141CrossRefGoogle Scholar
  11. R.M. Callaway and S.C. Pennings, Impact of a parasitic plant on the zonation of two salt marsh perennials. Oecologia 114 (1998) 100-105CrossRefGoogle Scholar
  12. P. Choler, R. Michalet and R.M. Callaway, Facilitation and competition on gradients in alpine plant communities. Ecology 82 (2001) 3295-3308CrossRefGoogle Scholar
  13. F.E. Clements, Plant Succession: An Analysis of the Development of Vegetation. Washington: Carnegie Institution of Washington edn (1916).Google Scholar
  14. J.H Connell and R.O. Slatyer, Mechanisms of succession in natural communities and their role in community stability and organization. Am. Nat. 111 (1977) 1119-1144CrossRefGoogle Scholar
  15. R. Moral del and A.F. Watson, Gradient structure of forest vegetation in the central Washington cascades. Vegetatio 38 (1978) 29-48CrossRefGoogle Scholar
  16. R.A. Dorich and D.W. Nelson, Ammonium measurement. Soil Sci. Soc. Am. J. 47 (1983) 833-836CrossRefGoogle Scholar
  17. H. Ellenberg, Physiologisches und ökologishes Verhalten derselben Pflanzenarten. Ber. Deutsch. Bot. Ges. 65 (1953) 351-362Google Scholar
  18. B. Finegan, Forest succession. Nature 312 (1984) 109-114CrossRefGoogle Scholar
  19. H.A. Gleason, The individualistic concept of the plant association. Bull. Torrey Bot. Club 53 (1926) 7-26CrossRefGoogle Scholar
  20. J.B. Grace, On the measurement of plant competition intensity. Ecology 76 (1995) 305-308CrossRefGoogle Scholar
  21. J.P. Grime, Vegetation classification by reference to strategies. Nature 250 (1974) 26-31CrossRefGoogle Scholar
  22. H.A.L. Henry and L.W. Aarssen, On the relationship between shade tolerance and shade avoidance strategies in woodland plants. Oïkos 80 (1997) 575-582Google Scholar
  23. M. Holmgren, M. Huston and M. Scheffer, The interplay of facilitation and competition in plant communities. Ecology 78 (1997) 1966-1975CrossRefGoogle Scholar
  24. S.H. Hurlbert, Pseudoreplication and the design of ecological field experiments. Ecol. Monogr. 54 (1984) 187-211CrossRefGoogle Scholar
  25. Joud D. 1997. Complexité morphogé né tique, historiqueet é cologique des ecosystems forestiers ‘hydromorphes’ des plateaux et terrasses du Bas–Dauphiné . PhD thesis, University of Grenoble.Google Scholar
  26. J.S. King, K.S. Pregitzer, D.R. Zak, M.E. Kubiske and W.E. Holmes, Correlation of foliage and litter chemistry of sugar mapleAcer saccharumas affected by elevated CO2 and varying N availability, and effects on decomposition. Oikos 94 (2001) 403-416CrossRefGoogle Scholar
  27. L.R. Lawlor, Direct and indirect effects of n-species competition. Oecologia 43 (1979) 355-364CrossRefGoogle Scholar
  28. J.M. Levine, Indirect facilitation: Evidence and predictions from a riparian community. Ecology 80 (1999) 1762-1769CrossRefGoogle Scholar
  29. S.H. Levine, Competitive interactions in ecosystems. Am. Nat. 110 (1976) 903-910CrossRefGoogle Scholar
  30. X. Li and S.D. Wilson, Facilitation among woody plants establishing in an old field. Ecology 79 (1998) 2694-2705CrossRefGoogle Scholar
  31. P. Liancourt, R.M. Callaway and R. Michalet, Stress tolerance and competitive response ability determine the outcome of biotic interactions. Ecology 86 (2004) 1611-1618Google Scholar
  32. T. Lippmaa, The unistratal concept of plant communities (The Unions). Am. Midland Nat. 21 (1939) 111-145CrossRefGoogle Scholar
  33. Ch.J. Lortie, R.W. Brooker, Ph. Choler, Z. Kikvidze, R. Michalet, F.I. Pugnaire and R.M. Callaway, Rethinking plant community theory. Oikos 107 (2004) 433-438CrossRefGoogle Scholar
  34. J.H. Markham and C.P. Chanway, Measuring plant neighbour effects. Funct. Ecol. 10 (1996) 548-549Google Scholar
  35. R.P. McIntosh, The continuum concept of vegetation. Bot. Rev. 33 (1967) 130-187Google Scholar
  36. R. Michalet and T. Philippe, Les Groupements acidiphiles (landes et pelouses) de l’étage subalpin des Monts Dore (Massif Central français): Syntaxonomie et potentialitiés dynamiques. Colloques Phytosociologiques 23 (1995) 433-471Google Scholar
  37. R. Michalet, C. Rolland, D. Joud, D. Gafta and R.M. Callaway, Associations between canopy and understory species increase along a rainshadow gradient in the Alps: habitat heterogeneity or facilitation?. Plant Ecol. 165 (2002) 145-160CrossRefGoogle Scholar
  38. T.E. Miller, Direct and indirect species interactions in an early old-field plant community. Am. Nat. 143 (1994) 1007-1025CrossRefGoogle Scholar
  39. D.J. Mladenoff, Dynamics of nitrogen mineralization and nitrification in hemlock and hardwood treefall gaps. Ecology 68 (1987) 1171-1180CrossRefGoogle Scholar
  40. E.H. Murchie and P. Horton, Contrasting patterns of photosynthetic acclimation to the light environment are dependent on the differential expression of the responses to altered irradiance and spectral quality. PlantCell Environ. 21 (1998) 203-214Google Scholar
  41. J.P. Pages and R. Michalet, A test of the indirect facilitation model in a temperate hardwood forest of the northern French Alps. J. Ecol. 91 (2003) 932-940CrossRefGoogle Scholar
  42. J.P. Pages, G. Pache, D. Joud, N. Magan and R. Michalet, Direct and indirect effects of shade on four forest tree seedlings in the French Alps. Ecology 84 (2003) 2741-2750Google Scholar
  43. D.A. Peltzer and M. Köchy, Competitive effects of grasses and woody plants in mixed-grass prairie. J. Ecol. 89 (2001) 519-527CrossRefGoogle Scholar
  44. S.C. Pennings and R.M. Callaway, Impact of a parasitic plant on the structure and dynamics of salt marsh vegetation. Ecology 77 (1996) 1410-1419CrossRefGoogle Scholar
  45. J.C. Rameau, D. Mansion and G. Dumé, Flore forestiére Française 2. France: Montagues, Institut pour Ie Développement Forestier (1993).Google Scholar
  46. G.H. Raschid and R. Schaeffer, Seasonal variation in the nitrogen mineralization and mineral nitrogen accumulation in two temperate forest soils. Pedobiologia 31 (1988) 381-390Google Scholar
  47. P.B. Reich, M.G. Tjoelker, M.B Walker, D.W. Vanderklein and C.C.C. Buschena, Close association of RGR, leaf and root morphology, seed mass and shade tolerance in seedling of nine boreal tree species grown in high and low light. Funct. Ecol. 12 (1998) 327-338CrossRefGoogle Scholar
  48. E.L. Rice and S.K. Pancholy, Inhibition of nitrification by climax ecosystems. II. Additional evidence and possible role of tannins. Am. J. Bot. 60 (1973) 691-702CrossRefGoogle Scholar
  49. L. Stone and A. Roberts, Conditions for a species to gain advantage from the presence of competitors. Ecology 72 (1991) 1964-1972CrossRefGoogle Scholar
  50. K. Taylor, A.P. Rowland and H.E. Jones, Molinia caerulea (L.) Moench. J. Ecol. 89 (2001) 126-144CrossRefGoogle Scholar
  51. K. Tielbörger and R. Kadmon, Indirect effects in a desert plant community: is competition among annuals more intense under shrub canopies?. Plant Ecol. 150 (2000) 53-63CrossRefGoogle Scholar
  52. D. Tilman, Resource Competition and Community Structure. New Jersey, U.S.A: Princeton (1982).Google Scholar
  53. J.H. Vandermeer, Indirect and diffuse interactions: complicated cycles in a population embedded in a large community. J. Theor. Biol. 142 (1990) 429-442Google Scholar
  54. J.F. Weltzin and G.R. Mcpherson, Faciliatation of conspecific seedling recruitment and shifts in temperate savanna ecotones. Ecol. Monogr. 69 (1999) 513-534CrossRefGoogle Scholar
  55. R.E. Wheatley, R. McDonald and A. McSmith, Extraction of nitrogen from soils. Biol. Fertility Soils 8 (1989) 189-190Google Scholar
  56. R.H. Whittaker, Vegetation of the Great Smoky Mountains. Ecol. Monogr. 26 (1956) 1-80CrossRefGoogle Scholar
  57. R.H. Whittaker, Vegetation of the Siskiyou moutains, Oregon and california. Ecol. Monogr. 30 (1960) 279-338CrossRefGoogle Scholar
  58. S.D. Wilson and P.A. Keddy, Species competitive ability and position along a natural stress/disturbance gradient. Ecology 67 (1986) 1236-1242CrossRefGoogle Scholar

Copyright information

© Springer 2006

Authors and Affiliations

  1. 1.CemagrefCampus universitaire Grenoble 1, UR Ecosystémes et Paysages MontagnardsSaint Martin d’HéresFrance
  2. 2.Laboratoire d’Ecologie des CommunautésUMR 1202 BIOGECO INRA-Université Bordeaux 1TalenceFrance

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