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

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.

[1]  C. Körner,et al.  Soil moisture effects determine CO2 responses of grassland species , 2000, Oecologia.

[2]  G. Bowes Facing the Inevitable: Plants and Increasing Atmospheric CO2 , 1993 .

[3]  C. Körner,et al.  In situ stomatal responses to long-term CO2 enrichment in calcareous grassland plants , 1997 .

[4]  J. Derner,et al.  Increasing CO 2 from subambient to superambient concentrations alters species composition and increases above-ground biomass in a C 3 / C 4 grassland , 2003 .

[5]  A. Knapp,et al.  9 – Ecosystem-Level Responses of Tallgrass Prairie to Elevated CO2 , 1996 .

[6]  S. Wand,et al.  Responses of wild C4 and C3 grass (Poaceae) species to elevated atmospheric CO2 concentration: a meta‐analytic test of current theories and perceptions , 1999 .

[7]  R. Sage Acclimation of photosynthesis to increasing atmospheric CO2: The gas exchange perspective , 1994, Photosynthesis Research.

[8]  B. D. Campbell,et al.  Elevated CO 2and water supply interactions in grasslands: a pastures and rangelands management perspective , 1997 .

[9]  C. Field,et al.  10 – Annual Grassland Responses to Elevated CO2 in Multiyear Community Microcosms , 1996 .

[10]  J. Coleman,et al.  Leaf conductance decreased under free-air CO2 enrichment (FACE) for three perennials in the Nevada desert , 2001 .

[11]  Responses in stomatal conductance to elevated CO2 in 12 grassland species that differ in growth form , 1996, Vegetatio.

[12]  A. Knapp,et al.  Biomass production and species composition change in a tallgrass prairie ecosystem after long‐term exposure to elevated atmospheric CO2 , 1999 .

[13]  J. Coleman,et al.  Photosynthetic down-regulation in Larrea tridentata exposed to elevated atmospheric CO2: Interaction with drought under glasshouse and field (FACE) exposure , 1998 .

[14]  W. Parton,et al.  Plant Nitrogen Dynamics in Shortgrass Steppe under Elevated Atmospheric Carbon Dioxide , 2004, Ecosystems.

[15]  P. Newton,et al.  Interaction of soil moisture and elevated CO2 on the above-ground growth rate, root length density and gas exchange of turves from temperate pasture , 1996 .

[16]  H. W. Hunt,et al.  Enhanced root system C-sink activity, water relations and aspects of nutrient acquisition in mycotrophic Bouteloua gracilis subjected to CO2 enrichment , 1994, Plant and Soil.

[17]  B. Drake,et al.  MORE EFFICIENT PLANTS: A Consequence of Rising Atmospheric CO2? , 1997, Annual review of plant physiology and plant molecular biology.

[18]  P. Newton,et al.  Photosynthetic responses of temperate species to free air CO2 enrichment (FACE) in a grazed New Zealand pasture , 2001 .

[19]  C. Körner,et al.  Carbon and water fluxes in a calcareous grassland under elevated CO2 , 1997 .

[20]  J. Coleman,et al.  Photosynthetic responses of Larrea tridentata to a step-increase in atmospheric CO2at the Nevada Desert FACE Facility , 2000 .

[21]  A. Knapp,et al.  Water vapour fluxes and their impact under elevated CO2 in a C4‐tallgrass prairie , 1997 .

[22]  C. Körner,et al.  A field study of the effects of elevated CO2 on plant biomass and community structure in a calcareous grassland , 1999, Oecologia.

[23]  M. Navas,et al.  Plant growth and competition at elevated CO2 : on winners, losers and functional groups. , 2003, The New phytologist.

[24]  A. Knapp,et al.  Effect of Elevated CO2 on Stomatal Density and Distribution in a C4 Grass and a C3 Forb under Field Conditions , 1994 .

[25]  J. Derner,et al.  Soil‐ and plant‐water dynamics in a C3/C4 grassland exposed to a subambient to superambient CO2 gradient , 2002 .

[26]  C. Körner,et al.  Biodiversity effects of elevated CO2 in species-rich model communities from the semi-arid Negev of Israel , 2001 .

[27]  R. B. Jackson,et al.  CO2 alters water use, carbon gain, and yield for the dominant species in a natural grassland , 1994, Oecologia.

[28]  R. B. Jackson,et al.  Nonlinear grassland responses to past and future atmospheric CO2 , 2002, Nature.

[29]  E. Kandeler,et al.  Six years of in situ CO2 enrichment evoke changes in soil structure and soil biota of nutrient‐poor grassland , 2003 .

[30]  C. Körner,et al.  The responses of alpine grassland to four seasons of CO2 enrichment: a synthesis , 1997 .

[31]  J. Roy,et al.  The influence of elevated CO2 on community structure, biomass and carbon balance of mediterranean old-fleld microcosms , 1995 .

[32]  R. B. Jackson,et al.  Stomatal acclimation over a subambient to elevated CO2 gradient in a C3/C4 grassland , 2002 .

[33]  P. Newton,et al.  Reduced water repellency of a grassland soil under elevated atmospheric CO2 , 2004 .

[34]  H. W. Polley,et al.  Elongated chambers for field studies across atmospheric CO2 gradients. , 2000 .

[35]  S. B. Idso,et al.  Increasing atmospheric CO2: effects on crop yield, water use and climate , 1983 .

[36]  W. Parton,et al.  Simulated Interaction of Carbon Dynamics and Nitrogen Trace Gas Fluxes Using the DAYCENT Model1 , 2006 .

[37]  C. Körner,et al.  Soil moisture dynamics of calcareous grassland under elevated CO2 , 1998, Oecologia.

[38]  Hendrik Poorter Interspecific variation in the growth response of plants to an elevated ambient CO2 concentration , 2004, Vegetatio.

[39]  J. Morgan,et al.  Soil and plant water relations determine photosynthetic responses of C3 and C4 grasses in a semi-arid ecosystem under elevated CO2. , 2003, Annals of botany.

[40]  F. A. Bazzaz,et al.  The Response of Natural Ecosystems to the Rising Global CO2 Levels , 1990 .

[41]  C. Körner,et al.  Seed production and seed quality in a calcareous grassland in elevated CO2 , 2003 .

[42]  A. Knapp,et al.  Photosynthetic Gas Exchange and Water Relation Responses of Three Tallgrass Prairie Species to Elevated Carbon Dioxide and Moderate Drought , 1997, International Journal of Plant Sciences.

[43]  J. Roy,et al.  9 – Responses to Elevated CO2 in Mediterranean Old-Field Microcosms: Species, Community, and Ecosystem Components , 1996 .

[44]  M. Lieffering,et al.  Increased Quantity and Quality of Coarse Soil Organic Matter Fraction at Elevated CO2 in a Grazed Grassland are a Consequence of Enhanced Root Growth Rate and Turnover , 2005, Plant and Soil.

[45]  B. D. Campbell,et al.  Effects of Elevated CO2 and Simulated Seasonal Changes in Temperature on the Species Composition and Growth Rates of Pasture Turves , 1994 .

[46]  Zong-ci Zhao,et al.  Climate change 2001, the scientific basis, chap. 8: model evaluation. Contribution of Working Group I to the Third Assessment Report of the Intergovernmental Panel on Climate Change IPCC , 2001 .

[47]  Alan K. Knapp,et al.  Photosynthetic and Water Relations Responses to Elevated CO2 in the C4 Grass Andropogon gerardii , 1993, International Journal of Plant Sciences.

[48]  Root dynamics in a semi-natural grassland in relation to atmospheric carbon dioxide enrichment, soil water and shoot biomass , 2000, Plant and Soil.

[49]  Christopher B. Field,et al.  Grassland responses to three years of elevated temperature, CO2, precipitation, and N deposition , 2003 .

[50]  C. Körner,et al.  Screen-aided CO2 control (SACC) : a middle ground between FACE and open-top chambers , 1997 .

[51]  H. W. Hunt,et al.  Photosynthetic pathway and ontogeny affect water relations and the impact of CO2 on Bouteloua gracilis (C4) and Pascopyrum smithii (C3) , 1998, Oecologia.

[52]  Jack A. Morgan,et al.  Consequences of growth at two carbon dioxide concentrations and two temperatures for leaf gas exchange in Pascopyrum smithii (C3) and Bouteloua gracilis (C4) , 1994 .

[53]  J. Ehleringer,et al.  Comparative ecophysiology of C3 and C4 plants , 1984 .

[54]  C. Körner,et al.  Growth and reproductive responses to elevated CO2 in wild cereals of the northern Negev of Israel , 2000 .

[55]  A. Knapp,et al.  Elevated CO2 and Leaf Longevity in the C4 Grassland‐Dominant Andropogon gerardii , 1999, International Journal of Plant Sciences.

[56]  H. W. Polley,et al.  Viewpoint: atmospheric CO2, soil water, and shrub/grass ratios on rangelands. , 1997 .

[57]  J. Roy,et al.  Short and long-term responses of whole-plant gas exchange to elevated CO2 in four herbaceous species , 2000 .

[58]  T. Mansfield,et al.  Nitrogen, phosphorus and potassium uptake and demand in Agrostis capillaris: the influence of elevated CO2 and nutrient supply. , 1995, The New phytologist.

[59]  J. Derner,et al.  Increasing CO2 from subambient to superambient concentrations alters species composition and increases above-ground biomass in a C3 /C4 grassland. , 2003, The New phytologist.

[60]  Jay M. Ham,et al.  Fluxes of CO2 and water vapor from a prairie ecosystem exposed to ambient and elevated atmospheric CO2 , 1995 .

[61]  Christian Körner,et al.  Biosphere responses to CO2 enrichment. , 2000 .

[62]  Craig T. Simmons,et al.  [The effect of climate]. , 2021, La Pathologie generale.

[63]  Mark Stitt,et al.  The interaction between elevated carbon dioxide and nitrogen nutrition: the physiological and molecular background , 1999 .

[64]  J. Morgan,et al.  Growth, gas exchange, leaf nitrogen and carbohydrate concentrations in NAD-ME and NADP-ME C4 grasses grown in elevated CO2 , 1998 .

[65]  R. B. Jackson,et al.  Gas exchange and photosynthetic acclimation over subambient to elevated CO2 in a C3–C4 grassland , 2001 .

[66]  Daniel G. Milchunas,et al.  CO2 ENHANCES PRODUCTIVITY, ALTERS SPECIES COMPOSITION, AND REDUCES DIGESTIBILITY OF SHORTGRASS STEPPE VEGETATION , 2004 .

[67]  E. Pendall,et al.  Partitioning evapotranspiration fluxes from a Colorado grassland using stable isotopes: Seasonal variations and ecosystem implications of elevated atmospheric CO2 , 2003, Plant and Soil.

[68]  J. Morgan,et al.  Elevated CO2 enhances water relations and productivity and affects gas exchange in C3 and C4 grasses of the Colorado shortgrass steppe. , 2001 .

[69]  B. D. Campbell,et al.  Crop ecosystem responses to climatic change: rangelands. , 2000 .

[70]  C. Field,et al.  CO2 effects on the water budget of grassland microcosm communities , 1997 .

[71]  J. Randerson,et al.  ELEVATED ATMOSPHERIC CO2 INCREASES WATER AVAILABILITY IN A WATER‐LIMITED GRASSLAND ECOSYSTEM 1 , 1997 .

[72]  T. Huxman,et al.  The effects of parental CO2 environment on seed quality and subsequent seedling performance in Bromusrubens , 1998, Oecologia.

[73]  D. Bremer,et al.  Effect of elevated atmospheric carbon dioxide and open-top chambers on transpiration in a tallgrass prairie , 1996 .

[74]  Jack A. Morgan,et al.  Responses of a C3 and a C4 perennial grass to elevated CO2 and temperature under different water regimes , 1996 .

[75]  B. Föger The FIELD study , 2006, The Lancet.

[76]  Alan K. Knapp,et al.  Biomass Production in a Tallgrass Prairie Ecosystem Exposed to Ambient and Elevated CO"2. , 1993, Ecological applications : a publication of the Ecological Society of America.

[77]  P. Reich,et al.  Leaf gas exchange responses of 13 prairie grassland species to elevated CO2 and increased nitrogen supply , 2001 .

[78]  J. Coleman,et al.  ELEVATED ATMOSPHERIC CO2 DOES NOT CONSERVE SOIL WATER IN THE MOJAVE DESERT , 2004 .

[79]  F. Chapin,et al.  Species‐specific responses of plant communities to altered carbon and nutrient availability , 2001 .

[80]  C. Körner,et al.  Long term effects of naturally elevated CO2 on mediterranean grassland and forest trees , 1994, Oecologia.

[81]  J. Morgan,et al.  Elevated CO2 increases soil moisture and enhances plant water relations in a long-term field study in semi-arid shortgrass steppe of Colorado , 2004, Plant and Soil.

[82]  A. Knapp,et al.  Elevated atmospheric CO2 alters stomatal responses to variable sunlight in a C4 grass , 1994 .

[83]  C. Körner,et al.  SYNTHESIS OF A SIX-YEAR STUDY OF CALCAREOUS GRASSLAND RESPONSES TO IN SITU CO2 ENRICHMENT , 2004 .

[84]  K. Knapp,et al.  Biomass production and species composition change in a tallgrass prairie ecosystem after long-term exposure to elevated atmospheric CO 2 , 1999 .

[85]  Dean L. Urban,et al.  Spatial Dependency of Vegetation–Environment Linkages in an Anthropogenically Influenced Wetland Ecosystem , 2004, Ecosystems.

[86]  C. Körner,et al.  Nutrient relations in calcareous grassland under elevated CO2 , 1998, Oecologia.

[87]  J. Coleman,et al.  Elevated CO2 increases productivity and invasive species success in an arid ecosystem , 2000, Nature.

[88]  T. Tschaplinski,et al.  Plant water relations at elevated CO2 -- implications for water-limited environments. , 2002, Plant, cell & environment.

[89]  R M Gifford,et al.  Stomatal sensitivity to carbon dioxide and humidity: a comparison of two c(3) and two c(4) grass species. , 1983, Plant physiology.

[90]  N. Stephenson Climatic Control of Vegetation Distribution: The Role of the Water Balance , 1990, The American Naturalist.

[91]  Effects of elevated CO2 (FACE) on the functional ecology of the drought-deciduous Mojave Desert shrub, Lycium andersonii , 2002 .

[92]  J. Houghton,et al.  Climate change 2001 : the scientific basis , 2001 .

[93]  P. Reich,et al.  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 , 2001 .

[94]  J. Coleman,et al.  Water use of two Mojave Desert shrubs under elevated CO2 , 2000 .

[95]  C. Körner,et al.  Differential phosphorus and nitrogen effects drive species and community responses to elevated CO2 in semi‐arid grassland , 2003 .

[96]  F. Miglietta,et al.  Elevated CO2 concentrations and stomatal density: observations from 17 plant species growing in a CO2 spring in central Italy , 1998 .

[97]  C. Owensby,et al.  The effect of CO2 enrichment on leaf photosynthetic rates and instantaneous water use efficiency of Andropogon gerardii in the tallgrass prairie , 2004, Photosynthesis Research.

[98]  Christopher B. Field,et al.  Stomatal responses to increased CO2: implications from the plant to the global scale , 1995 .

[99]  T. Huxman,et al.  Reproductive allocation and seed production in Bromus madritensis ssp. rubens at elevated atmospheric CO2 , 1999 .

[100]  T. Huxman,et al.  The Effects of Parental CO2 and Offspring Nutrient Environment on Initial Growth and Photosynthesis in an Annual Grass , 2001, International Journal of Plant Sciences.

[101]  J. Coleman,et al.  Biotic, abiotic and performance aspects of the Nevada Desert Free‐Air CO2 Enrichment (FACE) Facility , 1999 .

[102]  C. Körner,et al.  Interactive effects of elevated CO2, P availability and legume presence on calcareous grassland: results of a glasshouse experiment , 1999 .

[103]  W. Parton,et al.  DAYCENT and its land surface submodel: description and testing , 1998 .

[104]  P. Niklaus Effects of elevated atmospheric CO2 on soil microbiota in calcareous grassland , 1998 .

[105]  C. Körner,et al.  A LONG‐TERM FIELD STUDY ON BIODIVERSITY × ELEVATED CO2 INTERACTIONS IN GRASSLAND , 2001 .

[106]  Christopher B. Field,et al.  Grassland Responses to Global Environmental Changes Suppressed by Elevated CO2 , 2002, Science.