Department of Crop Sciences
Stephen Long
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Stephen Long

134 Institute for Genomic Biology
1206 W. Gregory Avenue
Urbana, IL 61801
(217) 333-2487


Ph.D.: Plant Sciences, Leeds University, England — 1976
B.Sc.: Agricultural Botany, Reading University, England — 1972

Research Areas

To understand the limitations to C4 photosynthesis and the adaptation of the process to cooler climates.
Advance the development of accessible mechanistic mathematical models relating environmental effects on photosynthesis to plant productivity (see:
Establish the potential of mitigation of atmospheric change through the development of herbaceous energy crops.
To understand crop responses to global atmospheric and climate change.
To understand mechanisms of plant responses to both rising atmospheric carbon dioxide concentration and tropospheric ozone, with particular reference to photosynthesis and relating changes at the molecular and biochemical level to observations of whole systems in the field.


Robert Emerson Professor of Crop Sciences and Plant Biology, University of Illinois, Champaign-Urbana. (1999 - present)
Professor (full) in Environmental Physiology and Head of Environment Biology Research Ground, University of Essex, England, UK. (1990 - 1998)
Reader in Environmental Physiology, University of Essex, England, UK. (1988 - 1990)
Senior Lecturer in Environmental Physiology, University of Essex, England, UK. (1987 - 1990)
Lecturer in Environmental Physiology, University of Essex, England, UK, (1975 - 1987, with tenure 1978)


Crop Sciences 431 - Plants and Global Change
Plant Biology 432 - Photosynthesis

Journal Articles

Bernacchi C.J., Kimball B.A., Quarles D.R., Long S.P. & Ort D.R. (2007) Decreases in stomatal conductance of soybean under open-air elevation of [CO2] are closely coupled with decreases in ecosystem evapotranspiration. Plant Physiology, 143, 134-144.
Long S.P., Ainsworth E.A., Leakey A.D.B., Ort D.R., Nosberger J. & Schimel D. (2007) Crop models, CO2, and climate change – Response. Science, 315, 460-460.
Wall G.W., Garcia R.L., Kimball B.A., Hunsaker D.J., Pinter P.J., Long S.P., Osborne C.P., Hendrix D.L., Wechsung F., Wechsung G., Leavitt S.W., LaMorte R.L. & Idso S.B. (2006) Interactive effects of elevated carbon dioxide and drought on wheat. Agronomy Journal, 98, 354-381.
Schroeder J.B., Gray M.E., Ratcliffe S.T., Estes R.E. & Long S.P. (2006) Effects of elevated Co-2 and O-3 on a variant of the western corn rootworm (Coleoptera: Chrysomelidae). Environmental Entomology, 35, 637-644.
Rogers A., Gibon Y., Stitt M., Morgan P.B., Bernacchi C.J., Ort D.R. & Long S.P. (2006) Increased C availability at elevated carbon dioxide concentration improves N assimilation in a legume. Plant Cell And Environment, 29, 1651-1658.
Bernacchi C.J., Leakey A.D.B., Heady L.E., Morgan P.B., Dohleman F.G., McGrath J.M., Gillespie K.M., Wittig V.E., Rogers A., Long S.P. & Ort D.R. (2006) Hourly and seasonal variation in photosynthesis and stomatal conductance of soybean grown at future CO2 and ozone concentrations for 3 years under fully open-air field conditions. Plant Cell And Environment, 29, 2077-2090.
Long S.P., Ainsworth E.A., Leakey A.D.B., Nosberger J. & Ort D.R. (2006) Food for thought: Lower-than-expected crop yield stimulation with rising CO2 concentrations. Science, 312, 1918-1921.
Morgan P.B., Mies T.A., Bollero G.A., Nelson R.L. & Long S.P. (2006) Season-long elevation of ozone concentration to projected 2050 levels under fully open-air conditions substantially decreases the growth and production of soybean. New Phytologist, 170, 333-343.
Leakey A.D.B., Bernacchi C.J., Ort D.R. & Long S.P. (2006) Long-term growth of soybean at elevated [CO2] does not cause acclimation of stomatal conductance under fully open-air conditions. Plant Cell And Environment, 29, 1794-1800.
Leakey A.D.B., Uribelarrea M., Ainsworth E.A., Naidu S.L., Rogers A., Ort D.R. & Long S.P. (2006) Photosynthesis, productivity, and yield of maize are not affected by open-air elevation of CO2 concentration in the absence of drought. Plant Physiology, 140, 779-790.
Farage P.K., Blowers D., Long S.P. & Baker N.R. (2006) Low growth temperatures modify the efficiency of light use by photosystem II for CO2 assimilation in leaves of two chilling-tolerant C-4 species, Cyperus longus L. and Miscanthus x giganteus. Plant Cell And Environment, 29, 720-728.
Dermody O., Long S.P. & DeLucia E.H. (2006) How does elevated CO2 or ozone affect the leaf-area index of soybean when applied independently? New Phytologist, 169, 145-155.
Davey P.A., Olcer H., Zakhleniuk O., Bernacchi C.J., Calfapietra C., Long S.P. & Raines C.A. (2006) Can fast-growing plantation trees escape biochemical down-regulation of photosynthesis when grown throughout their complete production cycle in the open air under elevated carbon dioxide? Plant Cell And Environment, 29, 1235-1244.
Long S.P., Zhu X.G., Naidu S.L. & Ort D.R. (2006) Can improvement in photosynthesis increase crop yields? Plant Cell And Environment, 29, 315-330.
Morgan P.B., Bollero G.A., Nelson R.L., Dohleman F.G. & Long S.P. (2005) Smaller than predicted increase in aboveground net primary production and yield of field-grown soybean under fully open-air [CO2] elevation. Global Change Biology, 11, 1856-1865.
Bernacchi C.J., Morgan P.B., Ort D.R. & Long S.P. (2005) The growth of soybean under free air [CO2] enrichment (FACE) stimulates photosynthesis while decreasing in vivo Rubisco capacity. Planta, 220, 434-446.
Ainsworth E.A. & Long S.P. (2005) What have we learned from 15 years of free-air CO2 enrichment (FACE)? A meta-analytic review of the responses of photosynthesis, canopy. New Phytologist, 165, 351-371.
Zhu X.G., Govindjee, Baker N.R., deSturler E., Ort D.R. & Long S.P. (2005) Chlorophyll a fluorescence induction kinetics in leaves predicted from a model describing each discrete step of excitation energy and electron transfer associated with photosystem II. Planta, 223, 114-133.
Pimentel C., Davey P.A., Juvik J.A. & Long S.P. (2005) Gene loci in maize influencing susceptibility to chilling dependent photoinhibition of photosynthesis. Photosynthesis Research, 85, 319-326.
Gielen B., Calfapietra C., Lukac M., Wittig V.E., De Angelis P., Janssens I.A., Moscatelli M.C., Grego S., Cotrufo M.F., Godbold D.L., Hoosbeek M.R., Long S.P., Miglietta F., Polle A., Bernacchi C.J., Davey P.A., Ceulemans R. & Scarascia-Mugnozza G.E. (2005) Net carbon storage in a poplar plantation (POPFACE) after three years of free-air CO2 enrichment. Tree Physiology, 25, 1399-1408.
Long S.P., Ainsworth E.A., Leakey A.D.B. & Morgan P.B. (2005) Global food insecurity. Treatment of major food crops with elevated carbon dioxide or ozone under large-scale fully open-air conditions suggests recent models may have overestimated future yields. Philosophical Transactions Of The Royal Society B-Biological Sciences, 360, 2011-2020.
Wittig V.E., Bernacchi C.J., Zhu X.G., Calfapietra C., Ceulemans R., Deangelis P., Gielen B., Miglietta F., Morgan P.B. & Long S.P. (2005) Gross primary production is stimulated for three Populus species grown under free-air CO2 enrichment from planting through canopy closure. Global Change Biology, 11, 644-656.
Davey P.A., Hunt S., Hymus G.J., DeLucia E.H., Drake B.G., Karnosky D.F. & Long S.P. (2004) Respiratory oxygen uptake is not decreased by an instantaneous elevation of [CO2], but is increased with long-term growth in the field at elevated [Co-2](1). Plant Physiology, 134, 520-527.
Zhu X.G., Portis A.R. & Long S.P. (2004) Would transformation of C-3 crop plants with foreign Rubisco increase productivity? A computational analysis extrapolating from kinetic properties to canopy photosynthesis. Plant Cell And Environment, 27, 155-165.
Leakey A.D.B., Bernacchi C.J., Dohleman F.G., Ort D.R. & Long S.P. (2004) Will photosynthesis of maize (Zea mays) in the US Corn Belt increase in future [CO2] rich atmospheres? An analysis of diurnal courses of CO2 uptake under free-air concentration enrichment (FACE). Global Change Biology, 10, 951-962.
Naidu S.L. & Long S.P. (2004) Potential mechanisms of low-temperature tolerance of C-4 photosynthesis in Miscanthus x giganteus: an in vivo analysis. Planta, 220, 145-155.
Rogers A., Allen D.J., Davey P.A., Morgan P.B., Ainsworth E.A., Bernacchi C.J., Cornic G., Dermody O., Dohleman F.G., Heaton E.A., Mahoney J., Zhu X.G., Delucia E.H., Ort D.R. & Long S.P. (2004) Leaf photosynthesis and carbohydrate dynamics of soybeans grown throughout their life-cycle under Free-Air Carbon dioxide Enrichment. Plant Cell And Environment, 27, 449-458.
Zhu X.G., Ort D.R., Whitmarsh J. & Long S.P. (2004) The slow reversibility of photosystem II thermal energy dissipation on transfer from high to low light may cause large losses in carbon gain by crop canopies: a theoretical analysis. Journal Of Experimental Botany, 55, 1167-1175.
Ainsworth E.A., Rogers A., Nelson R. & Long S.P. (2004) Testing the "source-sink" hypothesis of down-regulation of photosynthesis in elevated [CO2] in the field with single gene substitutions in Glycine max. Agricultural And Forest Meteorology, 122, 85-94.
Miyazaki S., Fredricksen M., Hollis K.C., Poroyko V., Shepley D., Galbraith D.W., Long S.P. & Bohnert H.J. (2004) Transcript expression profiles of Arabidopsis thaliana grown under controlled conditions and open-air elevated concentrations of CO2 and of O-3. Field Crops Research, 90, 47-59.
Morgan P.B., Bernacchi C.J., Ort D.R. & Long S.P. (2004) An in vivo analysis of the effect of season-long open-air elevation of ozone to anticipated 2050 levels on photosynthesis in soybean. Plant Physiology, 135, 2348-2357.
Long S.P., Ainsworth E.A., Rogers A. & Ort D.R. (2004) Rising atmospheric carbon dioxide: Plants face the future. Annual Review Of Plant Biology, 55, 591-628.
Heaton E., Voigt T. & Long S.P. (2004) A quantitative review comparing the yields of two candidate C-4 perennial biomass crops in relation to nitrogen, temperature and water. Biomass & Bioenergy, 27, 21-30.
Long S.P. & Bernacchi C.J. (2003) Gas exchange measurements, what can they tell us about the underlying limitations to photosynthesis? Procedures and sources of error. Journal Of Experimental Botany, 54, 2393-2401.
Morgan P.B., Ainsworth E.A. & Long S.P. (2003) How does elevated ozone impact soybean? A meta-analysis of photosynthesis, growth and yield. Plant Cell And Environment, 26, 1317-1328.
Naidu S.L., Moose S.P., Al-Shoaibi A.K., Raines C.A. & Long S.P. (2003) Cold tolerance of C-4 photosynthesis in Miscanthus x giganteus: Adaptation in amounts and sequence of C-4 photosynthetic enzymes. Plant Physiology, 132, 1688-1697.
Mason C.F., Underwood G.J.C., Baker N.R., Davey P.A., Davidson I., Hanlon G., Long S.P., Oxborough K., Paterson D.M. & Watson A. (2003) The role of herbicides in the erosion of salt marshes in eastern England. Environmental Pollution, 122, 41-49.
Bernacchi C.J., Calfapietra C., Davey P.A., Wittig V.E., Scarascia-Mugnozza G.E., Raines C.A. & Long S.P. (2003) Photosynthesis and stomatal conductance responses of poplars to free-air CO2 enrichment (PopFACE) during the first growth cycle and immediately following coppice. New Phytologist, 159, 609-621.
Ainsworth E.A., Tranel P.J., Drake B.G. & Long S.P. (2003) The clonal structure of Quercus geminata revealed by conserved microsatellite loci. Molecular Ecology, 12, 527-532.
Ainsworth E.A., Rogers A., Blum H., Nosberger J. & Long S.P. (2003) Variation in acclimation of photosynthesis in Trifolium repens after eight years of exposure to Free Air CO2 Enrichment (FACE). Journal Of Experimental Botany, 54, 2769-2774.
Bernacchi C.J., Pimentel C. & Long S.P. (2003) In vivo temperature response functions of parameters required to model RuBP-limited photosynthesis. Plant Cell And Environment, 26, 1419-1430.
Bernacchi C.J., Portis A.R., Nakano H., von Caemmerer S. & Long S.P. (2002) Temperature response of mesophyll conductance. Implications for the determination of Rubisco enzyme kinetics and for limitations to photosynthesis in vivo. Plant Physiology, 130, 1992-1998.
Ainsworth E.A., Davey P.A., Bernacchi C.J., Dermody O.C., Heaton E.A., Moore D.J., Morgan P.B., Naidu S.L., Ra H.S.Y., Zhu X.G., Curtis P.S. & Long S.P. (2002) A meta-analysis of elevated [CO2] effects on soybean (Glycine max) physiology, growth and yield. Global Change Biology, 8, 695-709.
Ainsworth E.A., Davey P.A., Hymus G.J., Drake B.G. & Long S.P. (2002) Long-term response of photosynthesis to elevated carbon dioxide in a Florida scrub-oak ecosystem. Ecological Applications, 12, 1267-1275.
McKee I.F. & Long S.P. (2001) Plant growth regulators control ozone damage to wheat yield. New Phytologist, 152, 41-51.
Hymus G.J., Dijkstra P., Baker N.R., Drake B.G. & Long S.P. (2001) Will rising CO2 protect plants from the midday sun? A study of photoinhibition of Quercus myrtifolia in a scrub-oak community in two seasons. Plant Cell And Environment, 24, 1361-1368.
Hymus G.J., Baker N.R. & Long S.P. (2001) Growth in elevated CO2 can both increase and decrease photochemistry and photoinhibition of photosynthesis in a predictable manner. Dactylis glomerata grown in two levels of nitrogen nutrition. Plant Physiology, 127, 1204-1211.
Geider R.J., Delucia E.H., Falkowski P.G., Finzi A.C., Grime J.P., Grace J., Kana T.M., La Roche J., Long S.P., Osborne B.A., Platt T., Prentice I.C., Raven J.A., Schlesinger W.H., Smetacek V., Stuart V., Sathyendranath S., Thomas R.B., Vogelmann T.C., Williams P. & Woodward F.I. (2001) Primary productivity of planet earth: biological determinants and physical constraints in terrestrial and aquatic habitats. Global Change Biology, 7, 849-882.
McLeod A.R. & Long S.P. (1999) Free-air carbon dioxide enrichment (FACE) in global change research: A review. In: Advances In Ecological Research, Vol 28, pp. 1-56.