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United States Department of Agriculture

Agricultural Research Service

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Prairie Heating and CO2 Enrichment (PHACE)

Mowll, W., D. M. Blumenthal, K. Cherwin, A. Smith, A. J. Symstad, L. T. Vermeire,  S. L. Collins, M. D. Smith.  2015.  Climatic controls of aboveground net primary production in semi‑arid grasslands along a latitudinal gradient portend low sensitivity to warming. Oecologia 177:959–969. DOI 10.1007/s00442-015-3232-7

Mueller, K. E., D. M. Blumenthal, Y. Carrillo, S. Cesarz, M. Ciobanu, J. Hines, S. Pabst, E. Pendall, C. M. de Tomasel , D. H. Wall, N. Eisenhauer. 2016. Elevated CO2 and warming shift the functional composition of soil nematode communities in a semiarid grassland. Soil Biology & Biochemistry 103:46-51

Ryan, E. M., K. Ogle, T.  J. Zelikova, D. R. Lecain, D. G. Williams, J. A. Morgan and E. Pendall , 2015. Antecedent moisture and temperature conditions modulate the response of ecosystem respiration to elevated CO2 and warming. Global Change Biology 21: 2588–2602, doi: 10.1111/gcb.12910

Mueller, K. E., D. M. Blumenthal, E. Pendall, Y. Carrillo, F. A. Dijkstra, D. G. Williams, R. F. Follett and J. A. Morgan. 2016. Impacts of warming and elevated CO2 on a semi-arid grassland are non-additive, shift with precipitation, and reverse over time. Ecology Letters, 19: 956–966

Chen, J.,  Y. Carrillo, E. Pendall, F. A. Dijkstra, R. D. Evans, J. A. Morgan, and D. G. Williams. 2015. Soil microbes compete strongly with plants for soil inorganic and amino acid nitrogen in a semiarid grassland exposed to elevated co2 and warming.  Ecosystems 18: 867–880, DOI: 10.1007/s10021-015-9868-7

Brzostek, E.R.,  J. M. Blair,  J. S . Dukes, S. D. Frey, S. E. Hobbie, J. M. Melillok, R. J. Mitchell , E. Pendall, P. B . Reich, G. R. Shaverk,  A. Stefanski, M. G. Tjoelker and A. C. Finzi. 2012. The effect of experimental warming and precipitation change on proteolytic enzyme activity: positive feedbacks to nitrogen availability are not universal. Global Change Biology 18:2617–2625, doi: 10.1111/j.1365-2486.2012.02685.x

Blumenthal, D.M., J.A. Kray, W. Ortmans, L.H. Ziska, E. Pendall. Cheatgrass is favored by warming but not CO2 enrichment in a semi-arid grassland.  Global Change Biology, doi: 10.1111/gcb.13278

Zelikova T.J., D.M .Blumenthal, D.G. Williams, L. Souza, D.R. LeCain, J.A. Morgan, E. Pendall. 2014. Long-term exposure to elevated CO2 enhances community stability by suppressing dominant plant species in a mixed-grass prairie. Proceedings of the National Academy of Sciences 111:15456-15461.

Reeves, J. L., D. M. Blumenthal, J. A. Kray, J. D. Derner. 2015. Increased seed consumption by biological control weevil tempers positive CO2 effect on invasive plant (Centaurea diffusa) fitness.Biological Control 84:36-43

LeCain D, D. Smith, J. Morgan, B.A. Kimball, E. Pendall, F. Miglietta. 2015. Microclimatic Performance of a Free-Air Warming and CO2 Enrichment Experiment in Windy Wyoming, USA. PLoS ONE 10(2): e0116834. doi:10.1371/journal.pone.0116834

Chen, J., T. J. Zelikova, E. Pendall,J. A. Morgan, D. G. Williams. 2014. Daily and seasonal changes in soil amino acid composition in a semiarid grassland exposed to elevated CO2 and warming. Biogeochemistry DOI 10.1007/s10533-014-0057-0

Suseela, V., D. Triebwasser-Freese, N. Linscheid, J. A. Morgan, and N. Tharayil. 2014. Litters of photosynthetically divergent grasses exhibit differential metabolic responses to warming and elevated CO2.Ecosphere

Reyes-Fox, M., H. Steltzer, M. J. Trlica, G. S. McMaster, A. A. Andales, D. R. LeCain & J. A. Morgan. 2014. Elevated CO2 further lengthens growing season under warming conditions. doi:10.1038/nature13207

Pendall, E.,  J. L. Heisler-White, D. G. Williams, F. A. Dijkstra, Y. Carrillo, J. A. Morgan, D. R. LeCain. 2013. Warming Reduces Carbon Losses from Grassland Exposed to Elevated Atmospheric Carbon Dioxide. 2013.. PLOSone august 2013: Vol. 8: issue 8.

Dijkstra, F. A., J. A. Morgan, R. F. Follett and D. R. LeCain. 2013. Climate change reduces the net sink of CH4 and N2O in a semi-arid grassland. Global Change Biology 19: 1816-1826. doi: 10.1111/gcb.12182.

Carrillo, Y.,  E. Pendall, F. A. Dijkstra, D. LeCain, J. A. Morgan, D. Blumenthal, S. Waldron. 2014. Disentangling root responses to climate change in a semiarid grassland. Oecologia, DOI 10.1007/s00442-014-2912-z.

Blumenthal, D. M., V. Resco, J. A. Morgan, D. G. Williams, D. R. LeCain, E. M. Hardy,E. Pendall and E. Bladyka. 2013. Invasive forb benefits from water savings by native plants and carbon fertilization under elevated CO2 and warming. New Phytologist, 200:1156-1165. doi: 10.1111/nph.12459.

Sorte, C.J.B., I. Ibanez, D.M. Blumenthal, N.A. Molinari, L.P. Miller, E.D. Grosholz, J.M. Diez, C.M. D'Antonio, J.D. Olden, S.J. Jones, and J.S. Dukes. Poised to prosper? A cross-system comparison of climate change effects on native and non-native species performance. Ecology Letters, 16:261-270, doi: 10.1111/ele.12017

Nie, M., E. Pendall, C. Bell, C. K. Gasch, S. Raut, S. Tamang, and M. D. Wallenstein. 2012. Positive climate feedbacks of soil microbial communities in a semiarid grassland.  Ecology Letters, 16:234-241. doi: 10.1111/ele.12034.

Tucker, C. L., J. Bell, E. Pendall, and K. Ogle.  2012. Does declining carbon-use efficiency explain thermal acclimation of soil respiration with warming?  Global Change Biology, 19:252-263, doi: 10.1111/gcb.12036.

Dijkstra, F. A., E. Pendall, J. A. Morgan, D. M. Blumenthal, Y. Carillo, D. R. LeCain, R. F. Follett, D. G. Williams. 2012. Climate change alters stoichiometry of phosphorus and nitrogen in semiarid grasslandNew Phytologist 196:807-815.

Dieleman,W .I .J., S .Vicca , F. A. Dijkstra, F. Hagedorn, M. J. Hovenden, K. S. Larsen, J. A. Morgan, A. Volder, C. S Beier, J. S. Dukes, J. King, S. Leuzinger, S. Linder, Y. Luo, R. Oren, P. DeAngelis, D. Tingey, M. R. Hoosbeek and I. A. Janssens. 2012.  Simple additive effects are rare: a quantitative review of plant biomass and soil process responses to combined manipulations of CO2 and temperature.Global Change Biology 18:2681-2693.

Carrillo, Y., F. A. Dijkstra, E. Pendall, J. A. Morgan, and D. M. Blumenthal. 2012. Controls over soil nitrogen pools in a semiarid grassland under elevated CO2 and warming.  Ecosystems 15:761-774.

Dijkstra, F. A., J. A. Morgan, J. C. von Fischer, and R. F. Follett. 2011. Elevated CO2 and warming effects on CH4 uptake in a semiarid grassland below optimum soil moisture. Journal Geophysical Research, 116, G01007, doi:10.1029/2010JG001288

Carrillo, Y., E. Pendall, F.A. Dijkstra, J.A. Morgan, and J.M. Newcomb.  2011.  Response of soil organic matter pools to elevated CO2 and warming in a semi-arid grasslandsPlant Soil 347:339-350.

Morgan, J. A., D. R. LeCain, E. Pendall, D. M. Blumenthal, B. A. Kimball, Y. Carrillo, D. G. Williams, J. Heisler-White, F. A. Dijkstra, and M. West. 2011.  C4 grasses prosper as carbon dioxide eliminates desiccation in warmed semi-arid grassland. Nature 476:202-206.

Dijkstra, F. A., D. Blumenthal, J. A. Morgan,E. Pendall, Y. Carrillo, and R. F. Follett. 2010. Contrasting effects of elevated CO2 and warming on
nitrogen cycling in a semiarid grassland.
New Phytologist 187: 426-437.

Bachman, S., J. L. Heisler-White, E. Pendall, D. G. Williams,
J.A. Morgan, and J. Newcomb. 2009. Elevated carbon dioxide alters impacts of precipitation pulses on ecosystem photosynthesis and respiration in a semi-arid
. Oecologia, 162:791-802.

Kimball, B. A., M. M. Conley, S. Wang, L. Xingwu, J. A. Morgan and D. P. Smith. 2008. Infra-red heater arrays for warming ecosystem field plots. Global Change Biology 14:309-320

Parton, W. J., J. A. Morgan, W. Guiming and S. J. Del Grosso. 2007. Projected Ecosystem Impact of the Prairie Heating and CO2 Enrichment Experiment.New Phytologist 174:823-834.


Selected papers from other Global Change Research, including Open Top Chamber Experiement (1997-2001) on Colorado Shortgrass Steppe


Dijkstra, F. A., J. A. Morgan, D. Blumenthal, R. F. Follett. 2010. Water limitation and plant inter-specific competition reduce rhizosphere-induced C decomposition and plant N uptake.Soil Biology and Biochemistry 42: 1073-1082.

Dijkstra,F. A., J. A. Morgan, D. R. LeCain, and R.F. Follett. 2010. Microbially mediated CH4 consumption and N2O emission is affected by elevated CO2, soil water content,and composition of semi-arid grassland species. Plant Soil  329:269-281.

Polley, H. W., W. Emmerich, J. A. Bradford, P. L. Sims, D. A. Johnson, N. Z. Saliendra, T. Svejcar, R. Angell, A. B. Frank, F. L. Phillips, K. A. Snyder, and J. A. Morgan. 2010. Physiological and environmental regulation of interannual variability in CO2 exchange on rangelands in the western United StatesGlobal Change Biology 16:990-1002.

Morgan, J. A., R. F. Follett, L. H. Allen, Jr., S. D. Grosso, J. D. Derner, F. Dijkstra, A. Franzluebbers, R. Fry, K. Paustian, and M. M. Schoeneberger.  2010.  Carbon sequestration in agricultural lands of the United States.  Journal of Soil and Water Conservation 65:6A-13A.  doi:10.2489/jswc.65.1.6A


Blumenthal, D., C. E. Mitchell, P. Py?ek, and V. Jaro??k.  2009.  Synergy between resource availability and pathogen release in plant invasion.  Proceedings of the National Academy of Sciences 106:7899-7904.

Shim, J. H., E. Pendall, J. A. Morgan, and D. S. Ojima. 2009. Wetting and drying cycles drive variations in the stable carbon isotope ratio of respired carbon dioxide in semi-arid grassland.  Oecologia doi 10.1007/s00442-009-1302-4.  

Alfieri, J., P. Blanken, J. A. Morgan,andD. P. Smith. 2009. Concerning the measurement and magnitude of heat, water vapor, and carbon dioxide exchange from a semiarid grassland. Journal of Applied Meteorology and Climatology 48:982-996.

Schuman, G. E., L. J. Ingram, P. D. Stahl, J. D. Derner, G. F. Vance, and J. A. Morgan. 2009. Influence of management on soil organic carbon dynamics in northern mixed-grass rangeland. In: R. Lal and R.F. Follett (eds.). Soil carbon sequestration and the greenhouse effect. Soil Science Society of America, Special Publication 57, second edition, Madison, WI. pp. 169-180.


Dijkstra, F. A, E. Pendall, A. Mosier, J. King, D. Milchunas, and J. A. Morgan. 2008. Long-term enhancement of N availability and plant growth under elevated CO2 in a semiarid grassland. Functional Ecology doi:10.1111/j.1365-2435.2008.01398.x.  

Svejcar, A. J., R. F. Angell, J. Bradford, W. Dugas, W. E. Emmerich, A. B. Frank, T. Gilmanov, M. R.  Haferkamp, D. A. Johnson, H. S. Mayeux Jr, P. Mielnick, J. A. Morgan, N. Saliendra, G. E. Schuman, P. L. Sims, K. A. Snyder. 2008. Carbon fluxes on North American rangelands. Rangeland Ecology & Management. 61:465-474.  

Ainsworth, E. A., C. Beier, C. Calfapietra, R. Cuelemans, M. Durand-Tardif, G. D. Farquhar, D. L.  Godbold, G. R. Hendrey, T. Hickler, J. Kaduk, D. J. Karnosky, B. A. Kimball, C. Koerner, M. Koornneef, T. Lafarge, A. D. B. Leakey, K. F. Lewin, S. P. Long, R. Manderscheid, D. L. McNeil, T. A. Meis, F. Miglietta, J. A. Morgan, J. Nagy, R. J. Norby, R. M. Norton, K. E. Percy, A. Rogers, J. Soussana, M. Stitt, H. Weigel, and J. W. White. 2008. Next generation of elevated [CO2] experiments with crops: A critical investment for feeding the future world. Plant Cell and Environment. 31:1317-1324.

Morgan, J. A., J. D. Derner, D. G. Milchunas and E. Pendall. Management implications of global change for Great Plains rangelands.  Rangelands.


Morgan, J. A., D. G. Milchunas, D. R. LeCain, M. S. West and A. Mosier. 2007. Carbon dioxide enrichment alters plant community structure and accelerates shrub growth in the shortgrass steppe. Proceedings of the Natl Academy of Sciences 104:14724-14729

K?rner, C., J. A. Morgan and R. Norby. 2007. CO2 fertilization: When, where, and how much? In: Canadell JG, Pataki DE, Pitelka LF (eds) Terrestrial Ecosystems in a Changing World. The IGBP Series. pp 9-22. Springer, Berlin.


LeCain, D. R., J. A. Morgan, D. G. Milchunas, A. R. Mosier, J. A. Nelson, and D. P. Smith. 2006. Root biomass of individual species, and root size characteristics after five years of CO2 enrichment on native shortgrass steppe.  Plant and Soil 279: 219-228


Derner, J. D., G. E. Schuman, M. Jawson, S. R. Shafer, J. A. Morgan, H. W. Polley, G. B. Runion, S. A. Prior, H. A. Torbert, H. H. Rogers, J. Bunce, L. Ziska, J. W. White, A. J. Franzluebbers, J. D. Reeder, R. Venterea and L .A. Harper. 2005.  Global Change Research on Rangelands and Pasturelands.Rangelands 27:36-42.

Pendall, E., J. Y. King, A. R. Mosier, J. Morgan and D. Milchunas. 2005. Stable isotope constraints on net ecosystem production under elevated CO2. In: L.B. Flanagan, J.R. Ehleringer and D.E. Pataki (eds) Stable Isotopes and Biosphere-Atmospheric Interactions: Processes and Biological Controls. pp 182-198. Elsevier Inc., San Diego, CA.

Milchunas, D. G., A. R. Mosier, J. A. Morgan, D. R. LeCain, J. Y. King,and J. A. Nelson. 2005. Elevated CO2 and defoliation effects on a shortgrass steppe: Forage quality versus quantity for ruminants.Agriculture, Ecosystems and Environment 111: 166-184.


Morgan, J. A., A. R. Mosier, D. G. Milchunas, D. R. LeCain, J. A. Nelson, and W. J. Parton. 2004. CO2 enhances productivity, alters species composition, and reduced digestibility of shortgrass steppe vegetation.Ecological Applications 14:208-219.

Morgan, J. A., D. E. Pataki, C. K?rner, H. Clark, S. J. Del Grosso, J. M. Gr?nzweig, 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 and M. R. Shaw. 2004. Water relations in grassland and desert ecosystems exposed to elevated atmospheric CO2.Oecologia 140:11-25.

Nelson, J. A., J. A. Morgan, D. R. LeCain, A. R. Mosier, D. G. Milchunas and W. J. Parton. 2004. Elevated CO2 increases soil moisture and enhances plant water relations in a long-term field study in the semi-arid shortgrass steppe of Northern Colorado. Plant and Soil 259:169-179.

Pendall, E. S., S. Bridgham, P. J. Hanson, B. Hungate, D. W. Kicklighter, D. W. Johnson, B. E. Law, Y. Luo, J. P. Megonigal, M. Olsrud, M. G. Ryan, P. Thornton and S. Wan. 2004. Belowground Process Responses to Elevated CO2 and Temperature: A Discussion of Observations, Measurement Methods, and Models.  New Phytologist 162  311-322 .

Pendall, E. l., J. Y. King, A. R. Mosier, J. Morgan and D. Milchunas.  2004.  Stable isotope constraints on net ecosystem production under elevated CO2.  IN L.B. Flanagan, J.R. Ehleringer and D.E. Pataki (eds) Stable Isotopes and Biosphere-Atmospheric Interactions: Processes and Biological Controls., pp 182-198,  Elsevier Inc., San Diego, CA.

Pendall, E., A. R. Mosier and J. A. Morgan. 2004. Rhizodeposition stimulated by elevated CO2 in a semi-arid grassland.New Phytologist 162: 447-458.


Mosier, A. R., E. Pendall and J. A. Morgan. 2003. Effect of water addition and nitrogen fertilization on the fluxes of CH4, CO2, NOx, and N2O following five years of elevated CO2 in the Colorado Shortgrass steppe.Atmospheric Chemistry and Physics Discussions 3:1-6.

Ferretti, D. F., E. Pendall, J. A. Morgan, J. A. Nelson, D. R. LeCain, A. R. Mostier. 2003. Partitioning evapotranspiration fluxes from a Colorado grassland using stable isotopes: Seasonal variations and ecosystem implications of elevated atmospheric CO2.Plant and Soil 254: 291-303.

LeCain, D. R., J. A. Morgan, A. R. Mosier and J. A. Nelson. 2003. Soil and plant water relations determine photosynthetic responses of C3 and C4 grasses in a semi-arid ecosystem under elevated CO2.Annals of Botany 92:41-52.

Pendall, E., S. Del Grosso, J. Y. King, D. R. LeCain, D. G. Milchunas, J. A. Morgan, A. R. Mosier, D. S. Ojima, W. A. Parton, P. P. Tans, and J. W. C. White. 2003. Elevated Atmospheric CO2 Effects and Soil Water Feedbacks on Soil Respiration Components in a Colorado grassland.Global Biogeochemical Cycles  17:1-13.


Morgan, J. A.  2002.  Looking beneath the surfaceScience 298:1903-1904.

Mosier, A. R., J. A. Morgan, J. Y. King, D. LeCain and D. G. Milchunas.  2002.  Soil-atmosphere exchange of CH4, CO2, NOx, and N2O in the Colorado Shortgrass Steppe under Elevated CO2.  Plant and Soil 240:201-211.


Morgan, J. A., D. R. LeCain, A. R. Mosier and D. G. Milchunas. 2001. Elevated CO2 enhances water relations and productivity and affects gas exchange in C3 and C4 grasses of the Colorado shortgrass steppe. Global Change Biology 7:451-466


Polley, H. W., J. A. Morgan, M. Stafford-Smith and B. Campbell.  2000.  Crop ecosystem responses to climatic change: rangelands, pp 293-314  IN K.R. Reddy and H.F. Hodges, eds, Climate Change and Global Crop Productivity, CAB International Publishing, UK. 



Last Modified: 2/3/2017
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