Plant and Soil

, Volume 224, Issue 2, pp 217–230 | Cite as

Plant diversity effects on soil heterotrophic activity in experimental grassland ecosystems

  • Eva M. Spehn
  • Jasmin Joshi
  • Bernhard Schmid
  • Jörn Alphei
  • Christian Körner


The loss of plant species from terrestrial ecosystems may cause changes in soil decomposer communities and in decomposition of organic material with potential further consequences for other ecosystem processes. This was tested in experimental communities of 1, 2, 4, 8, 32 plant species and of 1, 2 or 3 functional groups (grasses, legumes and non-leguminous forbs). As plant species richness was reduced from the highest species richness to monocultures, mean aboveground plant biomass decreased by 150%, but microbial biomass (measured by substrate induced respiration) decreased by only 15% (P = 0.05). Irrespective of plant species richness, the absence of legumes (across diversity levels) caused microbial biomass to decrease by 15% (P = 0.02). No effect of plant species richness or composition was detected on the microbial metabolic quotient (qCO2) and no plant species richness effect was found on feeding activity of the mesofauna (assessed with a bait-lamina-test). Decomposition of cellulose and birchwood sticks was also not affected by plant species richness, but when legumes were absent, cellulose samples were decomposed more slowly (16% in 1996, 27% in 1997, P = 0.006). A significant decrease in earthworm population density of 63% and in total earthworm biomass by 84% was the single most prominent response to the reduction of plant species richness, largely due to a 50% reduction in biomass of the dominant `anecic' earthworms. Voles (Arvicola terrestris L.) also had a clear preference for high-diversity plots. Soil moisture during the growing season was unaffected by plant species richness or the number of functional groups present. In contrast, soil temperature was 2 K higher in monocultures compared with the most diverse mixtures on a bright day at peak season. We conclude that the lower abundance and activity of decomposers with reduced plant species richness was related to altered substrate quantity, a signal which is not reflected in rates of decomposition of standard test material. The presence of nitrogen fixers seemed to be the most important component of the plant diversity manipulation for soil heterotrophs. The reduction in plant biomass due to the simulated loss of plant species had more pronounced effects on voles and earthworms than on microbes, suggesting that higher trophic levels are more strongly affected than lower trophic levels.

BIODEPTH decomposition earthworms legumes microbial biomass plant species richness 


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  1. Anderson J P E and Domsch K H 1978 A physiological method for the quantitative measurement of microbial biomass in soils. Soil Biol. Biochem. 10, 215–221.Google Scholar
  2. Bardgett R D and Shine A 1999 Linkages between plant litter diversity, soil microbial biomass and ecosystem function in temperate grasslands. Soil Biol. Biochem. 31, 317–321.Google Scholar
  3. Blair J M, Parmelee R W and Beare M H 1990 Decay rates, nitrogen fluxes, and decomposer communities of single-and mixed-species foliar litter. Ecology 71, 1976–1985.Google Scholar
  4. Bouché M B 1977 Ecologie et Paraécologie: Peut-on apprecier le role de la faune dans les cycles biogeochimiques. In Soil organisms as components of ecosystems. Eds U Lohm and T Persson. pp. 157–163. Ecological Bulletin, Stockholm.Google Scholar
  5. Chapman K, Whittaker J B and Heal O W 1988 Metabolic and faunal activity in litters of tree mixtures compared with pure stands. Agr. Ecosyst. Environ. 24, 33–40.Google Scholar
  6. Christie P, Newman EI and Campbell R, 1974 Grassland species can influence the abundance of microbes on each other's roots. Nature 250, 570–571.Google Scholar
  7. Christie P, Newman EI and Campbell R, 1978 The influence of neighbouring grassland plants on each other's endomycorrhizas and root surface microorganisms. Soil Biol. Biochem. 10, 521–527.Google Scholar
  8. Clarholm M 1985 Interactions of bacteria, protozoa and plants leading to mineralization of soil nitrogen. Soil Biol. Biochem. 17, 181–187.Google Scholar
  9. Corbet G B and Ovenden D 1982 Mammals of Britain and Europe. Paul Parey, Hamburg, Berlin.Google Scholar
  10. Darwin C 1881 The formation of vegetable mould through the action of worms. John Murray, London.Google Scholar
  11. Diemer M, Joshi J, Korner C, Schmid B and Spehn E 1997 An experimental protocol to assess the effects of plant diversity on ecosystem functioning utilized in a European research network. Bulletin of the Geobotanical Institute ETH 63, 95–107.Google Scholar
  12. Doak D F, Bigger D, Harding E K, Marvier M A, O'Malley R E and Thomson D 1998 The statistical inevitability of stabilitydiversity relationships in community ecology. Amer. Naturalist 151, 264–276.Google Scholar
  13. Dugas W A, Hicks R A and Gibbens R P 1996 Structure and function of C-3 and C-4 Chihuaihuan Desert plant communities: Energy balance components. J. Arid Environ. 34, 63–79.Google Scholar
  14. Edwards C A 1983 Earthworm ecology in cultivated soils. In Earthworm Ecology. From Darwin to vermiculture. Ed. J E Satchell. pp 123–138. Chapman & Hall, London.Google Scholar
  15. Edwards C A and Bohlen P J 1996 Biology and ecology of earthworms. Chapman & Hall, London.Google Scholar
  16. Edwards C A and Lofty J R 1972 Biology of Earthworms. Chapman & Hall. London.Google Scholar
  17. Edwards C A and Lofty J R 1978 The influence of arthropods and earthworms upon root growth of direct drilled cereals. J. Appl. Ecol. 17, 533–534.Google Scholar
  18. Edwards W M, Shipitalo M J, Owens L B and Norton L D 1990 Effect of Lumbricus terrestris L. burrows on hydrology of continuous no-till corn fields. Geoderma 46, 73–84.Google Scholar
  19. Ellenberg H 1988 Vegetation ecology of Central Europe. Cambridge University Press, Cambridge.Google Scholar
  20. Ewel J J, Mazzarino M J and Berish CW 1991 Tropical soil fertility changes under monocultures and successional communities of different structure. Ecol. Appl. 1, 289–302.Google Scholar
  21. Gaskin G J and Miller J D 1996 Measurement of soil water content using a simplified impedance measuring technique. J. Agric. Res. 63, 153–160.Google Scholar
  22. Griffiths B S, Welschen R, von Arendonk J J C M and Lambers H 1992 The effect of nitrate-nitrogen on bacteria and the bacterial feeding fauna in the rhizosphere of different grass species. Oecologia 91, 253–259.Google Scholar
  23. Groffman P M, Eagan P, Sullivan W M and Lemunyon J L 1996 Grass species and soil type effects on microbial biomass and activity. Plant Soil 183, 61–67.Google Scholar
  24. Hector A, Beale A J, Minns A, Otway S J and Lawton J H 2000 Consequences for the reduction of plant diversity for litter decomposition: effects through litter quality and microenvironment. Oikos, 90. (In press).Google Scholar
  25. Hector A, Schmid B, Beierkuhnlein C, Caldeira M C, Diemer M, Dimitrakopoulos P G, Finn J, Freitas H, Giller P S, Good J, Harris R, Högberg P, Huss-Danell K, Joshi J, Jumpponen A, Körner C, Leadley PW, Loreau M, Minns A, Mulder C P H, O'Donovan G, Otway S J, Pereira J S, Prinz A, Read D J, Scherer-Lorenzen M, Schulze E-D, Siamantziouras A-S D, Spehn E M, Terry A C, Troumbis A Y, Woodward F I, Yachi S and Lawton J H 1999 Plant diversity and productivity experiments in European grasslands. Science 286, 1123–1127.Google Scholar
  26. Holland E A and Coleman D C 1987 Litter placement effects on microbial and organic matter dynamics in an agroecosystem. Ecology 68, 425–433.Google Scholar
  27. Insam H and Domsch K H 1988 Relationship between soil organic carbon and microbial biomass on chronosequences of reclamation sites. Microbial Ecol. 15, 177-188.Google Scholar
  28. Jenkinson D S, Fox R H and Rayner J H 1985 Interactions between fertilizer nitrogen and soil nitrogen-the so-called ‘priming’ effect. J. Soil Sci. 36, 425–444.Google Scholar
  29. Joshi J, Matthies D and Schmid B 2000 Root hemiparasites and plant diversity in experimental grassland communities. J. Ecol. 88. (In press).Google Scholar
  30. Kelliher F M, Leuning R and Schulze E D 1993 Evaporation and canopy characteristics of coniferous forests and grasslands. Oecologia 95, 153–163.Google Scholar
  31. Lee K B 1985 Earthworms. Their ecology and relationships with soils and land use. Academic Press, Sydney.Google Scholar
  32. Leuning R, Condon A G, Dunin F X, Zegelin S and Denmead O T 1994 Rainfall interception and evaporation from soil below a wheat canopy. Agr. Forest Meteorol. 67, 221–238.Google Scholar
  33. McNaughton S J 1993 Biodiversity and function of grazing ecosystems. In Biodiversity and ecosystem function. Eds. E-D Schulze and H A Mooney. pp 361–383. Springer, Berlin.Google Scholar
  34. Mulder C P H, Koricheva J, Huss-Danell K, Hogberg P and Joshi J 1999 Insects affect relationships between plant species richness and ecosystem processes. Ecology Letters 2, 237–246.Google Scholar
  35. Naeem S, Thompson L J, Lawler S P, Lawton J H and Woodfin R M 1994 Declining biodiversity can alter the performance of ecosystems. Nature 368, 734–737.Google Scholar
  36. Naeem S, Thompson L J, Lawlers S P, Lawton J H and Woodfin R M 1995 Empirical evidence that declining species diversity may alter the performance of terrestrial ecosystems. Phil. Trans. Roy. Soc. London B 347, 249–262.Google Scholar
  37. Neter J and Wasserman W 1974 Applied Linear Statistical Models. Richard D. Irwin Inc., Homewood, Illinois.Google Scholar
  38. Niklaus P A 1998 Effects of elevated atmospheric CO2 on soil microbiota in calcareous grassland. Glob. Change Biol. 4, 451–458.Google Scholar
  39. Paul E A and Clark F E 1996 Soil microbiology and biochemistry. Academic Press, San Diego.Google Scholar
  40. Payne R W, Lane P W, Digby P G N, Harding S A, Leech P K, Morgan G W, Todd A D, Thompson R, Tunicliffe Wilson G, Welham S J and White R P 1993 GENSTAT 5 Reference Manual. Clarendon Press, Oxford.Google Scholar
  41. Ripley E A and Saugier B 1978 Biophysics of a natural grassland. J. Appl. Ecol. 15, 459–479.Google Scholar
  42. Roth C H and Joschko M 1991 A note on the reduction of runoff from crusted soils by earthworm burrows and artificial channels. Z. Pflanz. Bodenk. 154, 101–105.Google Scholar
  43. Rustad L 1994 Element dynamics along a decay continuum in a red spruce ecosystem in Maine, USA. Ecology 75, 867–879.Google Scholar
  44. Rychnovska M 1993 Structure and functioning of seminatural meadows. Developments in Agricultural and managed-forest Ecology 27, Elsevier, Amsterdam, London, NY.Google Scholar
  45. Scheu S 1992 Automated measurement of the respiratory response of soil microcompartments: Active microbial biomass in earthworm faeces. Soil Biol. Biochem. 24, 1113–1118.Google Scholar
  46. Schlesinger W H 1977 Carbon balance in terrestrial detritus. Annu. Rev. Ecol. Syst. 8, 51–81.Google Scholar
  47. Smith J and Paul E 1990 The significance of soil microbial biomass estimations. In Soil Biochemistry. Eds J Bollag and G Stotzky. pp 357–393. Marcel Dekker, New York.Google Scholar
  48. Spehn E M, Joshi J, Schmid B, Diemer M and Körner C 2000 Aboveground resource use increases with plant species richness in experimental grassland ecosystems. Funct. Ecol. 14. (In press).Google Scholar
  49. Staaf H 1980 lnfluence of chemical composition, addition of raspberry leaves, and nitrogen supply on decomposition rate and dynamics of nitrogen and phosphorus in beech leaf litter. Oikos 35, 55–62.Google Scholar
  50. Stephan A, Meyer AH and Schmid B 2000 Plant diversity affects culturable soil bacteria in experimental grassland communities. J. Ecol. (In press)Google Scholar
  51. Swift M J, Heal 0 W and Anderson J M 1979 Decomposition in terrestrial ecosystems. University of California Press, Berkeley, California, USA.Google Scholar
  52. Thielemann U 1986 Elektrischer Regenwurmfang mit der Oktett-Methode. Pedobiologia 29, 296–3O2.Google Scholar
  53. Thörne E 1990 Assessing feeding activities of soil-living animals. Bait-lamina-tests. Pedobiologia 34, 89–101.Google Scholar
  54. Tilman D, Wedin D and Knops J 1996 Productivity and sustainability influenced by biodiversity in grassland ecosystems. Nature 379, 718–720.Google Scholar
  55. Van de Geijn S C and van Veen J A 1993 Implications of increased carbon dioxide levels for carbon input and turnover in soils. Vegetatio 104/105, 283–292.Google Scholar
  56. Vossbrinck C R, Coleman D C and Woolley T A 1979 Abiotic and biotic factors in litter decomposition in a semi-arid grassland. Ecology 60, 265–271.Google Scholar
  57. Wardle D A, Bonner K I and Nicholson K 5 1997 Biodiversity and plant litter: Experimental evidence which does not support the view that enhanced species richness improves ecosystem function. Oikos 79, 247–258.Google Scholar
  58. Wardle D A and Nicholson K S 1996 Synergistic effects of grassland plant species on soil microbial biomass and activity: implications for ecosystem-level effects of enriched plant diversity. Funct. Ecol. 10, 410–416.Google Scholar
  59. Wedin D and Pastor J 1993 Nitrogen mineralisation dynamics in grass monocultures. Oecologia 96, 186–192.Google Scholar
  60. Wedin D A and Tilman D 1990 Species effects on nitrogen cycling: a test with perennial grasses. Oecologia 84, 433–441.Google Scholar
  61. Zak D R, Tilman D, Parmenter R R, Rice C W, Fisher F M, Vose J, Milchunas D and Martin C W 1994 Plant production and soil microorganisms in late-successional ecosystems: a continentalscale study. Ecology 75, 2333–2347.Google Scholar
  62. Zaller J G and Arnone J A 1999 Earthworm responses to plant species loss and elevated CO2 in calcareous grassland. Plant Soil 208, 1–8.Google Scholar
  63. Zaller J G and Arnone J A 1999 Interactions between plant species and earthworm casts in a calcareous grassland under elevated CO2. Ecology 80, 873–881.Google Scholar

Copyright information

© Kluwer Academic Publishers 2000

Authors and Affiliations

  • Eva M. Spehn
  • Jasmin Joshi
  • Bernhard Schmid
  • Jörn Alphei
  • Christian Körner

There are no affiliations available

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