, Volume 140, Issue 1, pp 11–25 | Cite as

Water relations in grassland and desert ecosystems exposed to elevated atmospheric CO2

  • J. A. Morgan
  • D. E. Pataki
  • C. Körner
  • H. Clark
  • S. J. Del Grosso
  • J. M. Grünzweig
  • A. K. Knapp
  • A. R. Mosier
  • P. C. D. Newton
  • P. A. Niklaus
  • J. B. Nippert
  • R. S. Nowak
  • W. J. Parton
  • H. W. Polley
  • M. R. Shaw
Concepts, Reviews, and Syntheses


Atmospheric CO2 enrichment may stimulate plant growth directly through (1) enhanced photosynthesis or indirectly, through (2) reduced plant water consumption and hence slower soil moisture depletion, or the combination of both. Herein we describe gas exchange, plant biomass and species responses of five native or semi-native temperate and Mediterranean grasslands and three semi-arid systems to CO2 enrichment, with an emphasis on water relations. Increasing CO2 led to decreased leaf conductance for water vapor, improved plant water status, altered seasonal evapotranspiration dynamics, and in most cases, periodic increases in soil water content. The extent, timing and duration of these responses varied among ecosystems, species and years. Across the grasslands of the Kansas tallgrass prairie, Colorado shortgrass steppe and Swiss calcareous grassland, increases in aboveground biomass from CO2 enrichment were relatively greater in dry years. In contrast, CO2-induced aboveground biomass increases in the Texas C3/C4 grassland and the New Zealand pasture seemed little or only marginally influenced by yearly variation in soil water, while plant growth in the Mojave Desert was stimulated by CO2 in a relatively wet year. Mediterranean grasslands sometimes failed to respond to CO2-related increased late-season water, whereas semiarid Negev grassland assemblages profited. Vegetative and reproductive responses to CO2 were highly varied among species and ecosystems, and did not generally follow any predictable pattern in regard to functional groups. Results suggest that the indirect effects of CO2 on plant and soil water relations may contribute substantially to experimentally induced CO2-effects, and also reflect local humidity conditions. For landscape scale predictions, this analysis calls for a clear distinction between biomass responses due to direct CO2 effects on photosynthesis and those indirect CO2 effects via soil moisture as documented here.


Biomass Carbon dioxide enrichment Landscape predictions Soil water Stomata 


  1. Adam NR, Owensby CE, Ham JM (2000) The effect of CO2 enrichment on leaf photosynthetic rates and instantaneous water use efficiency of Andropogon gerardii in the tallgrass prairie. Photosynth Res 65:121–129CrossRefGoogle Scholar
  2. Allard V, Newton PCD, Soussana J-F, Carran RA, Matthew C (2004) Increased quantity and quality of coarse soil organic matter fractions at elevated CO2 in a grazed grassland are a consequence of enhanced root growth and turnover. Plant Soil (in press)Google Scholar
  3. Anderson LJ, Maherali H, Johnson HB, Polley HW, Jackson RB (2001) Gas exchange and photosynthetic acclimation over subambient to elevated CO2 in a C3-C4 grassland. Global Change Biol 7:693–707CrossRefGoogle Scholar
  4. Bazzaz FA (1990) The response of natural ecosystems to the rising global CO2 levels. Annu Rev Ecol Syst 21:167–196Google Scholar
  5. Bettarini I, Vaccari F, Miglietta F (1998) Elevated CO2 concentrations and stomatal density: observations from 17 plant species growing in a CO2 spring in central Italy. Global Change Biol 4:17–22CrossRefGoogle Scholar
  6. Bowes G (1993) Facing the inevitable: plants and increasing atmospheric CO2. Annu Rev Plant Physiol Plant Mol Biol 44:309–332CrossRefGoogle Scholar
  7. Bremer DJ, Ham JM, Owensby CE (1996) Effect of elevated atmospheric carbon dioxide and open-top chambers on transpiration in a tallgrass prairie. J Environ Q 25:691–701Google Scholar
  8. Campbell BD, Stafford Smith DM, McKeon GM (1997) Elevated CO2 and water supply interactions in grasslands: a pastures and rangelands management perspective. Global Change Biol 3:177–187CrossRefGoogle Scholar
  9. Chiariello NR, Field CB (1996) Annual grassland responses to elevated CO2 in multiyear community microcosms. In: Körner C, Bazzaz FA (eds) Carbon dioxide, populations, and communities. Academic, San Diego, pp 140–157Google Scholar
  10. Del Grosso SJ, Parton WJ, Mosier AR, Hartman MD, Brenner J, Ojima DS, Schimel DS (2001) Simulated interaction of carbon dynamics and nitrogen trace gas fluxes using the DAYCENT model. In: Schaffer M, Ma L, Hansen S (eds) Modeling carbon and nitrogen dynamics for soil management. CRC, Boca Raton, Florida, pp 303–332Google Scholar
  11. Drake BG, Gonzàlez-Meler MA, Long SP (1997) More efficient plants: a consequence of rising atmospheric CO2. Annu Rev Plant Physiol Plant Mol Biol 48:609–639Google Scholar
  12. Edwards GR, Clark H, Newton PCD (2001) Carbon dioxide enrichment affects seedling recruitment in an infertile, permanent grassland grazed by sheep. Oecologia 127:383–394CrossRefGoogle Scholar
  13. Ferretti DF, Pendall E, Morgan JA, Nelson JA, LeCain DR, Mosier AR (2003) Partitioning evapotranspiration fluxes from a Colorado grassland using stable isotopes: seasonal variations and implication for elevated atmospheric CO2. Plant Soil 254:291–303CrossRefGoogle Scholar
  14. Field CB, Jackson RB, Mooney HA (1995) Stomatal responses to increased CO2: implications from the plant to the global scale. Plant Cell Environ 18:1214–1225Google Scholar
  15. Field CB, Lund CP, Chiariello NR, Mortimer BE (1997) CO2 effects on the water budget of grassland microcosm communities. Global Change Biol 3:197–206CrossRefGoogle Scholar
  16. Fredeen AL, Randerson JT, Holbrook NM, Field CB (1997) Elevated atmospheric CO2 increases water availability in a water-limited grassland ecosystem. J Am Water Resour Assoc 33:1033–1039Google Scholar
  17. Gill RA, Polley HW, Johnson HB, Anderson LJ, Maherali H, Jackson RB (2002) Nonlinear grassland responses to past and future atmospheric CO2. Nature 417:279–282CrossRefPubMedGoogle Scholar
  18. Grünzweig JM, Körner C (2000) Growth and reproductive responses to elevated CO2 in wild cereals of the northern Negev of Israel. Global Change Biol 6:231–238Google Scholar
  19. Grünzweig JM, Körner C (2001a) Growth, water and nitrogen relations in grassland model ecosystems of the semi-arid Negev of Israel exposed to elevated CO2. Oecologia 128:251–262CrossRefGoogle Scholar
  20. Grünzweig JM, Körner C (2001b) Biodiversity effects of elevated CO2 in species-rich model communities from the semi-arid Negev of Israel. Oikos 95:112–124Google Scholar
  21. Grünzweig JM, Körner C (2003) Differential P and N effects drive species and community responses to elevated CO2 in semi-arid grassland. Funct Ecol 17:766–777Google Scholar
  22. Ham JM, Owensby CE, Coyne PI, Bremer DJ (1995) Fluxes of CO2 and water vapor from a prairie ecosystem exposed to ambient and elevated atmospheric CO2. Agric For Meteorol 77:73–93CrossRefGoogle Scholar
  23. Hamerlynck EP, McCallister CA, Knapp AK, Ham JM, Owensby CE (1997) Photosynthetic gas exchange and water relation responses of three tallgrass prairie species to elevated carbon dioxide and moderate drought. Int J Plant Sci 158:608–616CrossRefGoogle Scholar
  24. Hamerlynck EP, Huxman TE, Nowak RS, Redar S, Loik M, Jordan DN, Zitzer SF, Coleman JS, Seemann JR, Smith SD (2000) Photosynthetic responses of Larrea tridentata to a step-increase in atmospheric CO2 at the Nevada Desert FACE facility. J Arid Environ 44:425–436CrossRefGoogle Scholar
  25. Hamerlynck EP, Huxman TE, Charlet TN, Smith SD (2002) Effects of elevated CO2 (FACE) on the functional ecology of the drought-deciduous Mojave Desert shrub, Lycium andersonii. Environ Exp Bot 48:93–106CrossRefGoogle Scholar
  26. Housman DC (2002) Effects of elevated CO2 on primary productivity in a Mojave Desert ecosystem. PhD Dissertation, University of Nevada, Las VegasGoogle Scholar
  27. Hunt HW, Elliott ET, Detling JK, Morgan JA, Chen D-X (1996) Responses of C3 and C4 perennial grass to elevated CO2 and climate change. Global Change Biol 2:35–47Google Scholar
  28. Huxman TE, Smith SD (2001) Photosynthesis in an invasive grass and native forb at elevated CO2 during an El Niño year in the Mojave Desert. Oecologia 128:193–201CrossRefGoogle Scholar
  29. Huxman TE, Hamerlynck EP, Jordan DN, Salsman KA, Smith SD (1998a) The effects of parental CO2 environment on seed quality and subsequent seedling performance in Bromus rubens. Oecologia 114:202–208CrossRefGoogle Scholar
  30. Huxman TE, Hamerlynck EP, Moore BD, Smith SD, Jordan DN, Zitzer SF, Nowak RS, Coleman JS, Seemann JR (1998b) Photosynthetic down-regulation in Larrea tridentata exposed to elevated atmospheric CO2: interaction with drought under glasshouse and field (FACE) exposure. Plant Cell Environ 21:1153–1161CrossRefGoogle Scholar
  31. Huxman TE, Hamerlynck EP, Smith SD (1999) Reproductive allocation and seed production in Bromus madritensis ssp. rubens at elevated CO2. Funct Ecol 13:769–777CrossRefGoogle Scholar
  32. Huxman TE, Charlet T, Grant C, Smith SD (2001) The effects of parental CO2 and offspring nutrient environment on initial growth and photosynthesis in an annual grass. Int J Plant Sci 162:617–623CrossRefGoogle Scholar
  33. IPCC, Working Group 1 Third Assessment Report (2001) Climate change 2001: the scientific basis. Cambridge University Press, Cambridge, UKGoogle Scholar
  34. Jackson RB, Sala OE, Field CB, Mooney HA (1994) CO2 alters water use, carbon gain, and yield for the dominant species in a natural grassland. Oecologia 98:257–262Google Scholar
  35. Joel G, Chapin FS III, Chiariello NR, Thayer SS, Field CB (2001) Species-specific responses of plant communities to altered carbon and nutrient availability. Global Change Biol 7:435–450CrossRefGoogle Scholar
  36. Johnson HB, Polley HW, Whitis RP (2000) Elongated chambers for field studies across atmospheric CO2 gradients. Funct Ecol 14:388–396CrossRefGoogle Scholar
  37. Jordan DN, Zitzer SF, Hendrey GR, Lewin KF, Nagy J, Nowak RS, Smith SD, Coleman JS, Seemann JR (1999) Biotic, abiotic and performance aspects of the Nevada Desert Free-Air CO2Enrichment (FACE) Facility. Global Change Biol 5:659–668CrossRefGoogle Scholar
  38. Kimball BA, Idso SB (1983) Increasing atmospheric CO2: effects on crop yield, water use and climate. Agric Water Manage 7:55–72Google Scholar
  39. King JY, Mosier AR, Morgan JA, LeCain DR, Milchunas DG, Parton WJ (2004) Plant nitrogen dynamics in shortgrass steppe under elevated atmospheric CO2. Ecosystems 7:147–160CrossRefGoogle Scholar
  40. Knapp AK, Hamerlynck EP, Owensby CE (1993) Photosynthetic and water relations responses to elevated CO2 in the C4 grass Andropogon gerardii. Int Plant Sci 154:459–466CrossRefGoogle Scholar
  41. Knapp AK, Cocke M, Hamerlynck EP, Owensby CE (1994a) Effect of elevated CO2 on stomatal density and distribution in a C4 grass and a C3 forb under field conditions. Ann Bot 74:595–599CrossRefGoogle Scholar
  42. Knapp AK, Fahnestock JT, Owensby CE (1994b) Elevated atmospheric CO2 alters stomatal responses to variable sunlight in a C4 grass. Plant Cell Environ 17:189–195Google Scholar
  43. Knapp AK, Hamerlynck EP, Ham JM, Owensby CE (1996) Responses in stomatal conductance to elevated CO2 in 12 grassland species that differ in growth form. Vegetatio 125:31–41Google Scholar
  44. Knapp AK, Bargmann N, Maragni LA, McAllister CA, Bremer DJ, Ham JM, Owensby CE (1999) Elevated CO2 and leaf longevity in the C4 grassland-dominant Andropogon gerardii. Int J Plant Sci 160:1057–1061CrossRefPubMedGoogle Scholar
  45. Körner C (2000) Biosphere responses to CO2 enrichment. Ecol Appl 10:1590–1619Google Scholar
  46. Körner C, Miglietta F (1994) Long term effects of naturally elevated CO2 on Mediterranean grassland and forest trees. Oecologia 99:343–351Google Scholar
  47. Körner C, Diemer M, Schäppi B, Niklaus P, Arnone J (1997) The responses of alpine grassland to four seasons of CO2 enrichment: a synthesis. Acta Oecol 18:165–175Google Scholar
  48. Lauber W, Körner C (1997) In situ stomatal responses to long-term CO2 enrichment in calcareous grassland plants. Acta Oecol 18:221–229Google Scholar
  49. Lauenroth WK, DG Milchunas (1991) The shortgrass steppe, Chapter 11. In: Coupland RT (ed) Natural grasslands: introduction and Western Hemisphere, Elsevier, New YorkGoogle Scholar
  50. Leadley PW, Niklaus P, Stocker R, Körner C (1997) Screen-aided CO2 control (SACC): a middle ground between FACE and open-top chambers. Acta Oecol 18:207–219Google Scholar
  51. Leadley PW, Niklaus PA, Stocker R, Körner C (1999) A field study of the effects of elevated CO2 on plant biomass and community structure in a calcareous grassland. Oecologia 118:39–49CrossRefGoogle Scholar
  52. LeCain DR, Morgan JA (1998) Growth, gas exchange, leaf nitrogen and carbohydrate concentrations in NAD-ME and NADP-ME C4 grasses grown in elevated CO2. Physiol Plant 102:297–306CrossRefGoogle Scholar
  53. LeCain DR, Morgan JA, Mosier AR, Nelson JA (2003) Soil and plant water relations, not photosynthetic pathway, primarily influence photosynthetic responses in a semi-arid ecosystem under elevated CO2. Ann Bot 92:41–52CrossRefGoogle Scholar
  54. Lee TD, Tjoelker NG, Ellsworth DS, Reich PB (2001) Leaf gas exchange responses of 13 prairie grassland species to elevated CO2 and increased nitrogen supply. New Phytol 150:405–418CrossRefGoogle Scholar
  55. Lund CP (2002) Ecosystem carbon and water budgets under elevated atmospheric carbon dioxide concentration in two California grasslands. Ph.D. thesis, Stanford University, StanfordGoogle Scholar
  56. Maherali H, Reid CD, Polley HW, Johnson HB, Jackson RB (2002) Stomatal acclimation over a subambient to elevated CO2 gradient in a C3/C4 grassland. Plant Cell Environ 25:557–566CrossRefGoogle Scholar
  57. Morgan JA, Hunt HW, Monz CA, LeCain DR (1994a) Consequences of growth at two carbon dioxide concentrations and two temperatures for leaf gas exchange in Pascopyrum smithii (C3) and Bouteloua gracilis (C4). Plant Cell Environ 17:1023–1033Google Scholar
  58. Morgan JA, Knight WG, Dudley LM, Hunt HW (1994b) Enhanced root system C-sink activity, water relations and aspects of nutrient acquisition in mycotrophic Bouteloua gracilis subjected to CO2 enrichment. Plant Soil 165:139–146Google Scholar
  59. Morgan JA, LeCain DR, Read JJ, Hunt HW, Knight WG (1998) Photosynthetic pathway and ontogeny affect water relations and the impact of CO2 on Bouteloua gracilis (C4) and Pascopyrum smithii (C3). Oecologia 114:483–493CrossRefGoogle Scholar
  60. Morgan JA, LeCain DR, Mosier AR, Milchunas DG (2001a) Elevated CO2 enhances water relations and productivity and affects gas exchange in C3 and C4 grasses of the Colorado shortgrass steppe. Global Change Biol 7:451–466CrossRefGoogle Scholar
  61. Morgan JA, Newton PCD, Nösberger J, Owensby CE (2001b) The influence of rising atmospheric CO2 on grassland ecosystems. Proceedings of the XIX International Grasslands Congress 2001. Brazilian Society of Animal Husbandry, Sao Pedro/Sao Paulo, Brazil, pp 973–980Google Scholar
  62. Morgan JA, Mosier AR, Milchunas DG, LeCain DR, Nelson JA, Parton WJ (2004) CO2 enhances productivity of the shortgrass steppe, alters species composition, and reduces forage digestibility. Ecol Appl 14:208–219Google Scholar
  63. Morrison JIL, Gifford RM (1983) Stomatal sensitivity to carbon dioxide and humidity. A comparison of two C3 and two C4 grass species. Plant Physiol 71:789–796Google Scholar
  64. Navas M-L, Guillerm J-L, Fabreguettes J, Roy J (1995) The influence of elevated CO2 on community structure, biomass and carbon balance of Mediterranean old-field microcosms. Global Change Biol 1:325–335Google Scholar
  65. Nelson JA, Morgan JA, LeCain DR, Mosier AR, Milchunas DG, Parton WJ (2004) Elevated CO2increases soil moisture and enhances plant water relations in a long-term field study in the semi-arid shortgrass steppe of Northern Colorado. Plant Soil 259:169–179CrossRefGoogle Scholar
  66. Newbery RM, Wolfenden J, Mansfield TA, Harrison AF (1995) Nitrogen, phosphorus and potassium uptake and demand in Agrostis capillaris: the influence of elevated CO2 and nutrient supply. New Phytol 130:565–574Google Scholar
  67. Newton PCD, Clark H, Bell CC, Glasgow EM, Campbell BD (1994) Effects of elevated CO2 and simulated seasonal changes in temperature on the species composition and growth rates of pasture turves. Ann Bot 73:53–59CrossRefGoogle Scholar
  68. Newton PCD, Clark H, Bell CC, Glasgow EM (1996) Interaction of soil moisture and elevated CO2 on the above-ground growth rate, root length density and gas exchange of turves from temperate pasture. J Exp Bot 47:771–779Google Scholar
  69. Newton PCD, Clark H, Edwards GR (2001) The effect of climate change on grazed grasslands. In: Shiyomi M, Koizumi H (eds) Structure and function of agroecosystem design and management. CRC, Boca Raton, Florida, pp 297–311Google Scholar
  70. Newton PCD, Carran RA, Lawrence EJ (2003) Reduced water repellency of a grassland soil under elevated atmospheric CO2. Global Change Biol 10:1–4Google Scholar
  71. Niklaus PA (1998) Effects of elevated atmospheric CO2 on soil microbiota in calcareous grasslands. Global Change Biol 4:451–458CrossRefGoogle Scholar
  72. Niklaus PA, Körner C (2004) Synthesis of a six year study of calcareous grassland responses to in situ CO2 enrichment. Ecol Monogr 74(3) (in press)Google Scholar
  73. Niklaus PA, Leadley PW, Stöcklin J, Körner C (1998a) Nutrient relations in calcareous grassland under elevated CO2. Oecologia 116:67–75CrossRefGoogle Scholar
  74. Niklaus PA, Spinnler D, Körner C (1998b) Soil moisture dynamics of calcareous grassland under elevated CO2. Oecologia 117:201–208CrossRefGoogle Scholar
  75. Niklaus PA, Leadley PW, Schmid B, Körner C (2001) A long-term field study on biodiversity × elevated CO2 interactions in grassland. Ecol Monogr 71:341–356Google Scholar
  76. Niklaus PA, Alphei J, Ebersberger D, Kandeler E, Tscherko D (2003) Six years of in situ CO2 enrichment evoke changes in soil structure and biota of nutrient-poor grassland. Global Change Biol 9:585–600Google Scholar
  77. Nowak RS, Jordan DN, DeFalco LA, Wilcox CS, Coleman JS, Seemann JR, Smith SD (2001) Leaf conductance decreased under free-air CO2 enrichment (FACE) for three perennials in the Nevada desert. New Phytol 150:449–458CrossRefGoogle Scholar
  78. Nowak RS, Zitzer SF, Babcock D, Smith-Longozo V, Charlet TN, Coleman JS, Seemann JR, Smith SD (2004) Elevated atmospheric CO2 does not conserve soil water in the Mojave Desert. Ecology 85:93–99Google Scholar
  79. Owensby CE, Coyne PI, Ham JM, Auen LM, Knapp AK (1993) Biomass production in a tallgrass prairie ecosystem exposed to ambient and elevated CO2. Ecol Appl 3:644–653Google Scholar
  80. Owensby CE, Ham JM, Knapp AK, Rice CW, Coyne PI, Auen LM (1996) Ecosystem level responses of tallgrass prairie to elevated CO2. In: Koch GW, Mooney HA (eds) Carbon dioxide and terrestrial ecosystems, Academic, San Diego, Calif., pp 147–162Google Scholar
  81. Owensby CE, Ham JM, Knapp AK, Bremer D, Auen LM (1997) Water vapour fluxes and their impact under elevated CO2 in a C4-tallgrass prairie. Global Change Biol 3:189–195CrossRefGoogle Scholar
  82. Owensby CE, Ham JM, Knapp AK, Auen LM (1999) Biomass production and species composition change in a tallgrass prairie ecosystem after long-term exposure to elevated atmospheric CO2. Global Change Biol 5:497–506CrossRefGoogle Scholar
  83. Parton WJ, Hartman MD, Ojima DS, Schimel DS (1998) DAYCENT: Its land surface submodel: description and testing. Global Planet Change 19:35–48CrossRefGoogle Scholar
  84. Pataki DE, Huxman TE, Jordan DN, Zitzer SF, Coleman JS, Smith SD, Nowak RS, Seemann JR (2000) Water use of two Mojave Desert shrubs under elevated CO2. Global Change Biol 6:889–898CrossRefGoogle Scholar
  85. Pearcy RW, Ehleringer J (1984) Comparative ecophysiology of C3 and C4plants. Plant Cell Environ 7:1–13Google Scholar
  86. Polley HW, Mayeux HS, Johnson HB, Tischler CR. (1997) Viewpoint: Atmospheric CO2, soil water, and shrub/grass ratios on rangelands. J. Range Manage 50:278–284Google Scholar
  87. Polley HW, Morgan JA, Campbell BD, Stafford Smith M (2000) Crop ecosystem responses to climatic change: rangelands. In: Reddy KR, Hodges HF (eds) Climate change and global crop productivity. CABI, Wallingford, Oxon, UK, pp 293–314Google Scholar
  88. Polley HW, Johnson HB, Derner JD (2002) Soil- and plant-water dynamics in a C3/C4 grassland exposed to a subambient to superambient CO2gradient. Global Change Biol 8:1118–1129CrossRefGoogle Scholar
  89. Polley HW, Johnson HB, Derner JD (2003) Increasing CO2 from subambient to superambient concentrations alters species composition and increases above-ground biomass in a C3/C4 grassland. New Phytol 160:319–327CrossRefGoogle Scholar
  90. Poorter J (1993) Interspecific variation in the growth response of plants to an elevated ambient CO2concentration. Vegetatio 104/105:77–97Google Scholar
  91. Poorter H, Navas M-L (2003) Plant growth and competition at elevated CO2; on winners, losers and functional groups. New Phytol 157:175–198CrossRefGoogle Scholar
  92. Reich PB, Tilman D, Craine J, Ellsworth D, Tjoelker MG, Knops J, Wedin D, Naeem S, Bahauddin D, Goth J, Bengtson W, Lee TD (2001) Do species and functional groups differ in acquisition and use of C, N and water under varying atmospheric CO2 and N availability regimes? A field test with 16 grassland species. New Phytol 150:435–448CrossRefGoogle Scholar
  93. Roumet C, Garnier E, Suzor H, Salager J-L, Roy J (2000) Short and long-term responses of whole-plant gas exchange to elevated CO2 in four herbaceous species. Environ Exp Bot 43:155–169CrossRefGoogle Scholar
  94. Roy J, Guillerm J-L, Navas M-L, Dhillion S (1996) Responses to elevated CO2 in Mediterranean old-field microcosms: species, community, and ecosystem components. In: Körner C, Bazzaz FA (eds) Carbon dioxide, populations, and communities. Academic, San Diego, Calif., pp 123–138Google Scholar
  95. Sage RF (1994) Acclimation of photosynthesis to increasing atmospheric CO2: the gas exchange perspective. Photosynth Res 39:351–368Google Scholar
  96. Shaw MR, Zavaleta ES, Chiariello NR, Cleland EE, Mooney HA, Field CB (2002) Grassland responses to global environmental changes suppressed by elevated CO2. Science 298:1987–1990CrossRefPubMedGoogle Scholar
  97. Sindhøj E, Hansson A-C, Andrén O, Kätterer T, Maarissink M, Peterssen R (2000) Root dynamics in a semi-natural grassland in relations to atmospheric carbon dioxide enrichment, soil water and shoot biomass. Plant Soil 223:253–263Google Scholar
  98. Smith SD, Huxman TE, Zitzer SF, Charlet TN, Housman DC, Coleman JS, Fenstermaker LK, Seemann JR, Nowak RS (2000) Elevated CO2 increases productivity and invasive species success in an arid ecosystem. Nature 408:79–82PubMedGoogle Scholar
  99. Stephenson NL (1990) Climatic control of vegetation distribution: the role of water balance. Am Nat 135:649–679CrossRefGoogle Scholar
  100. Stitt M, Krapp A (1999) The interaction between elevated carbon dioxide and nitrogen nutrition: the physiological and molecular background. Plant Cell Environ 22:583–621Google Scholar
  101. Stocker R, Leadley PW, Körner C (1997) Carbon and water fluxes in a calcareous grassland under elevated CO2. Funct Ecol 11:222–230Google Scholar
  102. Stöcklin J, Körner C (1998) Interactive effects of CO2, P availability and legume presence on calcareous grassland: results of a glasshouse experiment. Funct Ecol 13:200–209CrossRefGoogle Scholar
  103. Thürig B, Körner C, Stöcklin J (2003) Seed production and seed quality in a calcareous grassland in elevated CO2. Global Change Biol 9:873–884CrossRefGoogle Scholar
  104. Verville JH (2000) Biological N fixation in a California annual grassland: responses to elevated CO2 and effect on community N status and productivity. Ph.D. thesis, Stanford University, Stanford, Calif.Google Scholar
  105. Volk M, Niklaus PA, Körner C (2000) Soil moisture effects determine CO2 responses of grassland species. Oecologia 125:380–388CrossRefGoogle Scholar
  106. Von Caemmerer S, Ghannoum O, Conroy JP, Clark H, Newton PCD (2001) Photosynthetic responses of temperate species to free air CO2 enrichment (FACE) in a grazed New Zealand pasture. Aust J Plant Physiol 28:439–450Google Scholar
  107. Wand SJE, Midgley GF, Jones MH, Curtis PS (1999) Responses of wild C4 and C3 grass (Poaceae) species to elevated atmospheric CO2 concentrations: a meta-analytic test of current theories and perceptions. Global Change Biol 5:723–741CrossRefGoogle Scholar
  108. Wullschleger SD, Tschaplinski TJ, Norby RJ (2002) Plant water relations at elevated CO2—implications for water-limited environments. Plant Cell Environ 25:319–331CrossRefPubMedGoogle Scholar
  109. Zavaleta ES (2001) Influences of climate and atmospheric changes on plant diversity and ecosystem function in a California grassland. Ph.D. thesis, Stanford University, Stanford, Calif.Google Scholar
  110. Zavaleta ES, Shaw MR, Chiariello NR, Thomas BD, Cleland EE, Field CB, Mooney HA (2003) Responses of a California annual grassland community to three years of experimental climate change, elevated CO2, and N deposition. Ecological Monographs 73(4):585–604Google Scholar

Copyright information

© Springer-Verlag 2004

Authors and Affiliations

  • J. A. Morgan
    • 1
  • D. E. Pataki
    • 2
  • C. Körner
    • 3
  • H. Clark
    • 4
  • S. J. Del Grosso
    • 5
  • J. M. Grünzweig
    • 6
  • A. K. Knapp
    • 7
  • A. R. Mosier
    • 8
  • P. C. D. Newton
    • 4
  • P. A. Niklaus
    • 3
  • J. B. Nippert
    • 9
  • R. S. Nowak
    • 10
  • W. J. Parton
    • 5
  • H. W. Polley
    • 11
  • M. R. Shaw
    • 12
  1. 1.Rangeland Resources Research UnitUSDA Agricultural Research ServiceFort CollinsUSA
  2. 2.Dept. of BiologyUniversity of UtahSalt Lake CityUSA
  3. 3.Institute of BotanyUniversity of BaselBaselSwitzerland
  4. 4.AgResearchPalmerston NorthNew Zealand
  5. 5.Natural Resource Ecology LaboratoryColorado State UniversityFort CollinsUSA
  6. 6.Dept of Environmental Science and Energy ResearchWeizmann Institute of Science76100, RehovotIsrael
  7. 7.Department of BiologyColorado State UniversityFort CollinsUSA
  8. 8.Soil Plant Nutrient ResearchUSDA Agricultural Research ServiceFort CollinsUSA
  9. 9.Deptartment of BiologyKansas State UniversityManhattanUSA
  10. 10.Department NERS / MS 370University of NevadaRenoUSA
  11. 11.Grassland, Soil and Water Research LaboratoryUSDA Agricultural Research ServiceTempleUSA
  12. 12.California ChapterNature ConservancySan FranciscoUSA

Personalised recommendations