Plant and Soil

, Volume 288, Issue 1–2, pp 249–261 | Cite as

How do climate warming and plant species richness affect water use in experimental grasslands?

  • H. J. De Boeck
  • C. M. H. M. Lemmens
  • H. Bossuyt
  • S. Malchair
  • M. Carnol
  • R. Merckx
  • I. Nijs
  • R. Ceulemans
Original Paper


Climate warming and plant species richness loss have been the subject of numerous experiments, but studies on their combined impact are lacking. Here we studied how both warming and species richness loss affect water use in grasslands, while identifying interactions between these global changes. Experimental ecosystems containing one, three or nine grassland species from three functional groups were grown in 12 sunlit, climate-controlled chambers (2.25 m2 ground area) in Wilrijk, Belgium. Half of these chambers were exposed to ambient air temperatures (unheated), while the other half were warmed by 3°C (heated). Equal amounts of water were added to heated and unheated communities, so that warming would imply drier soils if evapotranspiration (ET) was higher. After an initial ET increase in response to warming, stomatal regulation and lower above-ground productivity resulted in ET values comparable with those recorded in the unheated communities. As a result of the decreased biomass production, water use efficiency (WUE) was reduced by warming. Higher complementarity and the improved competitive success of water-efficient species in mixtures led to an increased WUE in multi-species communities as compared to monocultures, regardless of the induced warming. However, since the WUE of individual species was affected in different ways by higher temperatures, compositional changes in mixtures seem likely under climatic change due to shifts in competitiveness. In conclusion, while increased complementarity and selection of water-efficient species ensured more efficient water use in mixtures than monocultures, global warming will likely decrease this WUE, and this may be most pronounced in species-rich communities.


Evapotranspiration Global warming Grassland species Species richness Water use efficiency 



Analysis of co-variance






General linear model

h (subscript)


mix (subscript)


mono (subscript)


res (subscript)



Species richness


Soil water content



u (subscript)



Water use efficiency



This research was funded by the Fund for Scientific Research—Flanders (Belgium) as project “Effects of biodiversity loss and climate warming on carbon sequestration mechanisms in terrestrial ecosystems”, contract # G.0434.03N. H.J. De Boeck holds a grant from the Institute for the Promotion of Innovation through Science and Technology in Flanders (IWT-Vlaanderen). H. Bossuyt is a post-doctoral research associate of the Fund for Scientific Research - Flanders. We thank B. Gielen for help with harvesting above-ground biomass, and F. Kockelbergh for technical assistance.


  1. Allen LH Jr, Pan D, Boote KJ, Pickering NB, Jones JW (2003) Carbon dioxide and temperature effects on evapotranspiration and water use efficiency of soybean. Agron J 95:1071–1081CrossRefGoogle Scholar
  2. Beyschlag W, Eckstein J (1998) Stomatal patchiness. Progr Bot 59:283–298Google Scholar
  3. Bruun HH, Ejrnæs R (2000) Classification of dry grassland vegetation in Denmark. J Veg Sci 11:585–596CrossRefGoogle Scholar
  4. Carter EB, Theodorou MK, Morris P (1997) Responses of Lotus corniculatus to environmental change. I. Effects of elevated CO2, temperature and drought on growth and plant development. New Phytol 136:245–253CrossRefGoogle Scholar
  5. Cernusca A (1976) Energy exchange within individual layers of a meadow. Oecologia 23:141–149CrossRefGoogle Scholar
  6. Chaves MM, Pereira JS, Maroco J, Rodrigues ML, Ricardo CPP, Osorio ML, Carvalho I, Faria T, Pinheiro C (2002) How plants cope with water stress in the field. Photosynthesis and growth. Ann Bot 89:907–916PubMedCrossRefGoogle Scholar
  7. Ciais P, Reichstein M, Viovy N, Granier A, Ogee J, Allard V, Aubinet M, Buchmann N, Bernhofer C, Carrara A, Chevallier F, De Noblet N, Friend AD, Friedlingstein P, Grunwald T, Heinesch B, Keronen P, Knohl A, Krinner G, Loustau D, Manca G, Matteucci G, Miglietta F, Ourcival JM, Papale D, Pilegaard K, Rambal S, Seufert G, Soussana JF, Sanz MJ, Schulze ED, Vesala T, Valentini R (2005) Europe-wide reduction in primary productivity caused by the heat and drought in 2003. Nature 437:529–533PubMedCrossRefGoogle Scholar
  8. De Boeck HJ, Nijs I, Lemmens CMHM, Ceulemans R (2006) Underlying effects of spatial aggregation (clumping) in relationships between plant diversity and resource uptake. Oikos 113:269–278CrossRefGoogle Scholar
  9. De Boeck HJ, Lemmens CMHM, Gielen B, Bossuyt H, Malchair S, Carnol M, Merckx R, Ceulemans R, Nijs I (2006) Combined effects of climate warming and plant diversity loss on above- and below-ground grassland productivity. Environ Exp Bot (in press)Google Scholar
  10. Eatherall A (1997) Modelling climate change impacts on ecosystems using linked models and a GIS. Clim Change 35:17–34CrossRefGoogle Scholar
  11. Ejrnæs R, Bruun HH (2000) Gradient analysis of dry grassland vegetation in Denmark. J Veg Sci 11:573–584CrossRefGoogle Scholar
  12. Gielen B, De Boeck H, Lemmens CMHM, Valcke R, Nijs I, Ceulemans R (2005) Grassland species will not necessarily benefit from future elevated air temperatures: a chlorophyll fluorescence approach to study autumn physiology. Physiol Plant 125:52–63CrossRefGoogle Scholar
  13. Haith DA, Shoemaker LL (1987) Generalized watershed loading functions for stream-nutrients. Water Resour Bull 23:471–478Google Scholar
  14. Hooper DU, Chapin FS, Ewel JJ, Hector A, Inchausti P, Lavorel S, Lawton JH, Lodge DM, Loreau M, Naeem S, Schmid B, Setala H, Symstad AJ, Vandermeer J, Wardle DA (2005) Effects of biodiversity on ecosystem functioning: a consensus of current knowledge. Ecol Monogr 75:3–35Google Scholar
  15. Houghton JT, Ding Y, Griggs DJ, Noguer M, van der Linden PJ, Dai X, Maskell K, Johnson CA (2001) IPCC 2001. Climate change 2001: the scientific basis. Contribution of working group I to the second assessment report of the Intergovernmental Panel on Climate Change. Cambridge University Press, CambridgeGoogle Scholar
  16. Larcher W (2003) Physiological plant ecology, 4th edn. Springer-Verlag, BerlinGoogle Scholar
  17. Lemmens CMHM, De Boeck HJ, Gielen B, Bossuyt H, Malchair S, Carnol M, Merckx R, Nijs I, Ceulemans R (2006) End-of-season effects of elevated temperature on ecophysiological processes of grassland species at different species richness levels. Environ Exp Bot 56:245–254CrossRefGoogle Scholar
  18. Llorens L, Penuelas J, Estiarte M (2003) Ecophysiological responses of two Mediterranean shrubs, Erica multiflora and Globularia alypum, to experimental drier and warmer conditions. Physiol Plant 119:231–243CrossRefGoogle Scholar
  19. Loreau M, Hector A (2001) Partitioning selection and complementarity in biodiversity experiments. Nature 412:72–76PubMedCrossRefGoogle Scholar
  20. Mahmoud A, Grime JP (1976) An analysis of competitive ability in three perennial grasses. New Phytol 77:431–435CrossRefGoogle Scholar
  21. Marchand FL, Nijs I, De Boeck HJ, Kockelbergh F, Mertens S, Beyens L (2004) Increased turnover but little change in the carbon balance of high-arctic tundra exposed to whole growing season warming. Arc Antarct Alp Res 36:298–307CrossRefGoogle Scholar
  22. Myneni RB, Keeling CD, Tucker CJ, Asrar G, Namani RR (1997) Increased plant growth in the northern high latitudes from 1981 to 1991. Nature 386:698–702CrossRefGoogle Scholar
  23. Naeem S, Thompson LJ, Lawler SP, Lawton JH, Woodfin RM (1994) Declining biodiversity can alter the performance of ecosystems. Nature 368:734–737CrossRefGoogle Scholar
  24. Naeem S, Li SB (1997) Biodiversity enhances ecosystem reliability. Nature 390:507–509CrossRefGoogle Scholar
  25. Reichstein M, Tenhunen JD, Roupsard O, Ourcival JM, Rambal S, Miglietta F, Peressotti A, Pecchiari M, Tirone G, Valentini R (2002) Severe drought effects on ecosystem CO2 and H2O fluxes at three Mediterranean evergreen sites: revision of current hypotheses? Global Change Biol 8:999–1017CrossRefGoogle Scholar
  26. Rodrigues ML, Pacheco CMA, Chaves MM (1995) Soil–plant water relations, root distribution and biomass partitioning in Lupinus albus L. under drought conditions. J Exp Bot 46:947–956Google Scholar
  27. Sala OE, Chapin FS III, Armesto JJ, Berlow E, Bloomfield J, Dirzo R, Huber-Sanwald E, Huenneke LF, Jackson RB, Kinzig A, Leemans R, Lodge DM, Mooney HA, Oesterheld M, Le Roy Poff N, Sykes MT, Walker BH, Walker M, Wall DH (2000) Global bio-diversity scenarios for the year 2100. Science 287:1770–1774PubMedCrossRefGoogle Scholar
  28. Saleska SR, Harte J, Torn MS (1999) The effect of experimental ecosystem warming on CO2 fluxes in a montane meadow. Global Change Biol 5:125–141CrossRefGoogle Scholar
  29. Thornley JHM, Cannell MGR (1997) Temperate grassland responses to climate change: an analysis using the Hurley pasture model. Ann Bot 80:205–221CrossRefGoogle Scholar
  30. Van Peer L, Nijs I, Reheul D, De Cauwer B (2004) Species richness and susceptibility to heat and drought extremes in synthesized grassland ecosystems: compositional vs physiological effects. Funct Ecol 18:769–778CrossRefGoogle Scholar
  31. van Ruijven J, Berendse F (2005). Diversity-productivity relationships: initial effects, long-term patterns, and underlying mechanisms. Proc Nat Sci USA 102:695–700CrossRefGoogle Scholar
  32. Walther G-R (2003) Plants in a warmer world. Perspect Plant Ecol Evol Syst 6:169–185CrossRefGoogle Scholar
  33. Xu ZZ, Zhou GS (2005) Effects of water stress and nocturnal temperature on carbon allocation in the perennial grass, Leymus chinensis. Physiol Plant 123:272–280CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2006

Authors and Affiliations

  • H. J. De Boeck
    • 1
  • C. M. H. M. Lemmens
    • 1
  • H. Bossuyt
    • 2
  • S. Malchair
    • 3
  • M. Carnol
    • 3
  • R. Merckx
    • 2
  • I. Nijs
    • 1
  • R. Ceulemans
    • 1
  1. 1.Research Group of Plant and Vegetation Ecology, Department of BiologyUniversity of Antwerp (Campus Drie Eiken)WilrijkBelgium
  2. 2.Division Soil and Water Management, Faculty of Bioscience EngineeringKatholieke Universiteit LeuvenLeuven/HeverleeBelgium
  3. 3.Laboratory of Plant and Microbial Ecology, Institute of Plant Biology B22University of LiègeLiègeBelgium

Personalised recommendations