Responses of a C 3 and C 4 perennial grass to C O 2 enrichment and climate change : Comparison between model predictions and experimental data

Ecological responses to C O 2 enrichment and climate change are expressed at several interacting levels: photosynthesis and stomatal movement at the leaf level, energy and gas exchanges at the canopy level, photosynthate allocation and plant growth at the plant level, and water budget and nitrogen cycling at the ecosystem level. Predictions of these ecosystem responses require coupling of ecophysiological and ecosystem processes. Version GEM2 of the grassland ecosystem model linked biochemical, ecophysiological and ecosystem processes in a hierarchical approach. The model included biochemical level mechanisms of C 3 and C4 photosynthetic pathways to represent direct effects of CO 2 on plant growth, mechanistically simulated biophysical processes which control interactions between the ecosystem and the atmosphere, and linked with detailed biogeochemical process submodels. The model was tested using two-year full factorial (CO2, temperature and precipitation) growth chamber data for the grasses Pascopyrum smithii (C 3) and Bouteloua gracilis (C4). The C3-C 4 photosynthesis submodels fitted the measured photosynthesis data from both the C 3 and the C 4 species subjected to different CO2, temperature and precipitation conditions. The whole GEM2 model accurately fitted plant biomass dynamics and plant N content data over a wide range of temperature, precipitation and atmospheric CO 2 concentration. Both data and simulation results showed that elevated CO 2 enhanced plant biomass production in both P. smithii (C 3) and B. gracilis (C4). The enhancement of shoot production by elevated CO 2 varied with temperature and precipitation. Doubling CO 2 increased modeled annual net primary production (NPP) of P. smithii by 36% and 43% under normal and elevated temperature regimes, respectively, and increased NPP of B. gracilis by 29% and 24%. Doubling CO 2 decreased modeled net N mineralization rate (N min) of soil associated with P. smithii by 3% and 2% at normal and high temperatures, respectively. N min of B. gracilis soil decreased with doubled CO 2 by 5% and 6% at normal and high temperatures. NPP increased with precipitation. The average NPP and N min of P. smithii across the treatments was greater than that of B. gracilis. In the C 3 species the response of NPP to increased temperatures was negative under dry conditions with ambient CO2, but was positive under wet conditions or doubled CO 2. The * Corresponding author. 0304-3800/96/$15.00 © 1996 Elsevier Science B.V. All rights reserved SSDI 0304-3800(94)00199-5 12 D.-X. Chen et al. / Ecological Modelling 87 (1996) 11-27 responses of NPP to elevated C O 2 in the C 4 species were positive under all temperature and precipitation treatments. N min increased with precipitation in both the C 3 and C 4 species. Elevated CO 2 decreased N min in the C4 system. The effects of elevated CO 2 on N_min in the C 3 system varied with precipitation and temperature. Elevated temperature decreased N rain under dry conditions, but increased it under wet conditions. Thus, there are strong interactions among the ef~cts of CO 2 enrichment, precipitation, temperature and species on NPP and N min. Interactions between ecophysiological processes and ecosystem processes were strong. GEM2 coupled these processes, and was able to represent the interactions and feedbacks that mediate ecological responses to CO 2 enrichment and climate change. More information about the feedbacks between water and N cycling is required to further validate the model. More experimental and modeling efforts are needed to address the possible effects of CO 2 enrichment and climate change on the competitive balance between different species in a plant community and the feedbacks to ecosystem function.

[1]  F. Bazzaz,et al.  Using growth analysis to interpret competition between a C3 and a C4 annual under ambient and elevated CO2 , 1989, Oecologia.

[2]  H. W. Polley,et al.  Increasing CO2 and plant-plant interactions: effects on natural vegetation , 2004, Vegetatio.

[3]  William K. Lauenroth,et al.  A gap dynamics simulation model of succession in a semiarid grassland , 1990 .

[4]  John Pastor,et al.  State-of-the-Art of Models of Production-Decomposition Linkages in Conifer and Grassland Ecosystems. , 1991, Ecological applications : a publication of the Ecological Society of America.

[5]  J. Reynolds,et al.  Long-Term Response of an Arctic Sedge to Climate Change: A Simulation Study. , 1992, Ecological applications : a publication of the Ecological Society of America.

[6]  M. Trlica,et al.  Interacting Effects of Soil Water, Temperature and Irradiance on CO 2 Exchange Rates of Two Dominant Grasses of the Shortgrass Prairie , 1977 .

[7]  Michael B. Coughenour,et al.  Mathematical simulation of C4 grass photosynthesis in ambient and elevated CO2 , 1994 .

[8]  G. Collatz,et al.  Coupled Photosynthesis-Stomatal Conductance Model for Leaves of C4 Plants , 1992 .

[9]  S. Idso,et al.  Tree growth in carbon dioxide enriched air and its implications for global carbon cycling and maximum levels of atmospheric CO2 , 1993 .

[10]  D. Jordan,et al.  The CO2/O2 specificity of ribulose 1,5-bisphosphate carboxylase/oxygenase , 1984, Planta.

[11]  John L. Monteith,et al.  A four-layer model for the heat budget of homogeneous land surfaces , 1988 .

[12]  Gordon B. Bonan Do biophysics and physiology matter in ecosystem models? , 1993 .

[13]  W. Parton,et al.  Analysis of factors controlling soil organic matter levels in Great Plains grasslands , 1987 .

[14]  M. Coughenour A mechanistic simulation analysis of water use, leaf angles, and grazing in East African graminoids , 1984 .

[15]  R. Monson,et al.  Photosynthetic adaptation to temperature in four species from the Colorado shortgrass steppe: a physiological model for coexistence , 1983, Oecologia.

[16]  W. Pockman,et al.  Interactions between C3 and C4 salt marsh plant species during four years of exposure to elevated atmospheric CO2 , 2004, Vegetatio.

[17]  F. Bazzaz,et al.  Terrestrial Plant Communities , 2019, CO2 and Plants.

[18]  P. Curtis,et al.  Elevated atmospheric CO2 and feedback between carbon and nitrogen cycles , 1993, Plant and Soil.

[19]  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.

[20]  Derek Eamus,et al.  The interaction of rising CO2 and temperatures with water use efficiency , 1991 .

[21]  F. S. Nakayama,et al.  Effects of increasing atmospheric CO2 on vegetation , 2004, Vegetatio.

[22]  J. Berry,et al.  A biochemical model of photosynthetic CO2 assimilation in leaves of C3 species , 1980, Planta.

[23]  I. E. Woodrow,et al.  A Model Predicting Stomatal Conductance and its Contribution to the Control of Photosynthesis under Different Environmental Conditions , 1987 .

[24]  W. Oechel,et al.  Carbon balance in tussock tundra under ambient and elevated atmospheric CO2 , 1990, Oecologia.

[25]  Donald A. Klein,et al.  Simulation model for the effects of climate change on temperate grassland ecosystems , 1991 .

[26]  B. Drake A field study of the effects of elevated CO2 on ecosystem processes in a Chesapeake Bay Wetland , 1992 .

[27]  De Chen,et al.  GEMTM: a general model for energy and mass transfer of land surfaces and its application at the FIFE sites , 1994 .

[28]  B. Legg Principles of Environmental Physics (second edition). By J. L. Monteith and M. H. Unsworth. Sevenoaks, Kent: Edward Arnold (1990), pp. 291, £14.95, hardback £30.00. , 1990, Experimental Agriculture.

[29]  I. R. Johnson,et al.  Temperature Dependence of Plant and Crop Process , 1985 .

[30]  De Chen,et al.  Interactive effects of CO2 enrichment and temperature on the growth of dioecious Hydrilla verticillata , 1994 .

[31]  N. Sionit,et al.  Environmental Controls on the Growth and Yield of Okra. I. Effects of Temperature and CO2 Enrichment at Cool Temperature1 , 1981 .

[32]  A. Dalcher,et al.  A Simple Biosphere Model (SIB) for Use within General Circulation Models , 1986 .

[33]  J. Goudriaan,et al.  Crop Micrometeorology: A Simulation Study , 1977 .

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

[35]  J. Coleman,et al.  Effects of CO_2 and Temperature on Growth and Resource Use of Co‐Occurring C_3 and C_4 Annuals , 1992 .

[36]  H. Shugart A Theory of Forest Dynamics , 1984 .

[37]  G. Hornberger,et al.  Empirical equations for some soil hydraulic properties , 1978 .

[38]  E. Rastetter,et al.  Potential Net Primary Productivity in South America: Application of a Global Model. , 1991, Ecological applications : a publication of the Ecological Society of America.

[39]  Stephen P. Long,et al.  Modification of the response of photosynthetic productivity to rising temperature by atmospheric CO2 concentrations: Has its importance been underestimated? , 1991 .