Professor George Ratcliffe

Professor of Plant Sciences, Head of Department

Tel +44 (0) 1865 275001
Fax +44 (0) 1865 275074

Research Area

Structure and function of the plant metabolic network.

Research Description

Metabolic networks supply the precursors, energy and reducing power required for the synthesis and turnover of cellular components. The associated flows of material – the metabolic fluxes – are crucial in determining the performance and productivity of cells and organisms. For example, in an agricultural context, the production of harvestable end-products of plant metabolism is entirely dependent on the flux phenotype of the plant; while in biotechnology, the exploitation of micro-organisms and plants hinges on an ability to reconfigure the metabolic network to favour a flux distribution that leads to the preferential synthesis of particular products. Thus the fluxes supported by the plant metabolic network play a pivotal role in determining both phenotype and productivity.

My main interest lies in understanding the organisation and regulation of the metabolic fluxes that occur in the plant metabolic network. A knowledge of the transcriptome, proteome or metabolome does not lead easily to the metabolic flux phenotype, and internal fluxes within the metabolic network have to be deduced from a suite of computational and experimental tools. My research group is strongly involved in the development and application of steady-state metabolic flux analysis (MFA), a technique that allows fluxes to be deduced from a stoichiometric model of the network using stable isotope (13C) labelling data and measurements of biosynthetic outputs. We complement this MFA work with an in silico approach using genome-scale models and constraints-based flux balance analysis. Together these methods allow us to assess the metabolic phenotypes of wild type, mutant and transgenic plants, and thus the metabolic impact of genetic and environmental perturbations.

Future work will address fundamental questions relating to metabolic network capacity and performance in the tissues of C3, C4 and CAM plants; as well as analyzing the metabolic interactions that occur between cell types in plant tissues, and between plants and microbes.

Current projects include:

  • Integrated source-sink metabolic modelling of tomato fruit development.
  • Constraints-based metabolic modelling of C4 and CAM photosynthesis.
  • A flux analysis perspective on the metabolic capacity of Euglena gracilis.
  • Flux analysis of Azorhizobium caulinodans under N2 and non-N2 fixing conditions.
  • Effect of charge and proton balancing on constraints-based metabolic models.
  • Metabolic analysis of potassium stress in an Arabidopsis suspension culture.

Other areas of interest for future projects:

  • Cell- and tissue-specific metabolic flux analysis.
  • In silico engineering of CAM photosynthesis and bioenergy crops.
  • Provision of reducing power for growth and development.
  • Metabolic analysis of osmotolerance in plant pathogens.
  • Metabolic flux analysis of symbiotic nitrogen fixation.

Publications

Recent publications and papers in press

M.T. Rossi, M. Kalde, C. Srisakvarakul, N.J. Kruger and R.G. Ratcliffe (2017) Cell-type specific metabolic flux analysis: a challenge for metabolic phenotyping and a potential solution in plants. Metabolites 7, 59. doi:10.3390/metabo704005.

N.J. Kruger, S.K. Masakapalli and R.G. Ratcliffe (2017) Assessing metabolic flux in plants with radiorespirometry. In Plant Respiration and Internal Oxygen: Methods and Protocols (ed K.J. Gupta) Methods in Molecular Biology 1670. 1-16.

Y. Zhang, K.F.M. Beard, C. Swart, S. Bergmann, I. Krahnert, Z. Nikoloski, A. Graf, R.G. Ratcliffe, L.J. Sweetlove, A.R. Fernie and T. Obata (2017) Protein-protein interactions and metabolite channelling in the plant tricarboxylic acid cycle. Nature Communications 8, 15212.

K.J. Gupta, C.P. Lee and R.G. Ratcliffe (2017) Nitrite protects mitochondrial structure and function under hypoxia. Plant and Cell Physiology 58, 175-183.

S.L. McCraw, D.H. Park, R. Jones, M. Bentley, A. Rico, R.G. Ratcliffe, N.J. Kruger, A. Collmer and G.M. Preston (2016) GABA uptake via the GABA permease GabP represses virulence gene expression in Pseudomonas syringae pv. tomato DC3000. Molecular Plant-Microbe Interactions 29, 938-949.

J.J. Terpolilli, S.K. Masakapalli, R. Karunakaran, I. Webb, R. Green, N.J. Watmough, N.J. Kruger, R.G. Ratcliffe and P.S. Poole (2016) Lipogenesis and redox balance in nitrogen-fixing pea bacteroids. Journal of Bacteriology 198, 2864-2875.

C.Y.M. Cheung, R.G. Ratcliffe and L.J. Sweetlove (2015) A method of accounting for enzyme costs in flux balance analysis reveals alternative pathways and metabolite stores in an illuminated Arabidopsis leaf. Plant Physiology 169, 1671-1682.

B. Royo, J.F. Morán, R.G. Ratcliffe and K.J. Gupta (2015) Nitric oxide induces the alternative oxidase pathway in Arabidopsis seedlings deprived of inorganic phosphate. Journal of Experimental Botany 66, 6273-6280.

N.J. Kruger and R.G. Ratcliffe (2015) Fluxes through plant metabolic networks: measurements, predictions, insights and challenges. Biochemical Journal 465, 27-38.

A.U. Igamberdiev, R.G. Ratcliffe and K.J. Gupta (2014) Plant mitochondria: source and target for nitric oxide. Mitochondrion 19, 329-333.

K.J. Gupta, L.A.J. Mur and R.G. Ratcliffe (2014) Guarding the guard cells? New Phytologist 203, 349-351.

S.K. Masakapalli, N.J. Kruger and R.G. Ratcliffe (2014) The metabolic flux phenotype of heterotrophic Arabidopsis cells reveals a flexible balance between the cytosolic and plastidic contributions to carbohydrate oxidation in response to phosphate limitation. The Plant Journal 78, 964-977.

C.Y.M. Cheung, M.G. Poolman, D.A. Fell, R.G. Ratcliffe and L.J. Sweetlove (2014) A diel flux-balance model captures interactions between light and dark metabolism during day-night cycles in C3 and CAM leaves. Plant Physiology 165, 917-929.

B.L. Møller and R.G. Ratcliffe (2014) Synthetic plant biology: the roots of a bio-based society. Current Opinion in Biotechnology 26, ix-xvi.

K.J. Gupta, K.H. Hebelstrup, N.J. Kruger and R.G. Ratcliffe (2014) Nitric oxide is required for homeostasis of oxygen and reactive oxygen species in barley roots under aerobic conditions. Molecular Plant 7, 747-750.

S.K. Masakapalli, A. Ritala, L. Dong, A.R. van der Krol, K.M. Oksman-Caldentey, R.G. Ratcliffe and L.J. Sweetlove (2014) Metabolic flux phenotype of tobacco hairy roots engineered for increased geraniol production. Phytochemistry 99, 73-85.

S.K. Masakapalli, R.G. Ratcliffe and T.C.R. Williams (2014) Quantification of 13C enrichments and isotopomer abundances for metabolic flux analysis using 1D NMR spectroscopy. In Plant Metabolic Flux Analysis: Methods and Protocols (eds M. Dieuaide-Noubhani and A.P. Alonso), Methods in Molecular Biology 1090, 73-86.

N.J. Kruger, S.K. Masakapalli and R.G. Ratcliffe (2014) Optimisation of steady-state 13C-labelling experiments for metabolic flux analysis. In Plant Metabolic Flux Analysis: Methods and Protocols (eds M. Dieuaide-Noubhani and A.P. Alonso), Methods in Molecular Biology 1090, 53-72.

杨亦扬,马立锋,黎星辉,R.G.Ratcliffe,N.J. Kruger, 阮建云 (2013) 氮素水平对茶树新梢叶片代谢谱及其昼夜变化的影响. 茶叶科学 33, 491-499.

C.Y.M. Cheung, T.C.R. Williams, M.G. Poolman, D.A. Fell, R.G. Ratcliffe and L.J. Sweetlove (2013) A method for accounting for maintenance costs in flux balance analysis improves the prediction of plant cell metabolic phenotypes under stress conditions. The Plant Journal 75, 1050-1061.

L.J. Sweetlove, T.C.R. Williams, C.Y.M. Cheung and R.G. Ratcliffe (2013) Modelling metabolic CO2 evolution – a fresh perspective on respiration. Plant, Cell and Environment 36, 1631-1640.

S.K. Masakapalli, N.J. Kruger and R.G. Ratcliffe (2013) The metabolic flux phenotype of heterotrophic Arabidopsis cells reveals a complex response to changes in nitrogen supply. The Plant Journal 74, 569-582.

J. Gerendas and R.G. Ratcliffe (2013) Root pH regulation. In Plant Roots: The Hidden Half, 4th edn (eds A. Eshel and T. Beeckman), Chapter 23, pp 1-18; CRC Press, Boca Raton, USA.

M.J. Morgan, S. Osorio, B. Gehl, C.J. Baxter, N.J. Kruger, R.G. Ratcliffe, A.R. Fernie and L.J. Sweetlove (2013) Metabolic engineering of tomato fruit organic acid content guided by biochemical analysis of an introgression line. Plant Physiology 161, 397-407.

Y.Y. Yang, X.H. Li, R.G. Ratcliffe and J.Y. Ruan (2013) Characterization of ammonium and nitrate uptake and assimilation in roots of tea plants. Russian Journal of Plant Physiology 60, 91-99.

M. Wirtz, K.F.M. Beard, C.P. Lee, A. Boltz, M. Schwarzländer, C. Fuchs, A.J. Meyer, C. Heeg, L.J. Sweetlove, R.G. Ratcliffe and R. Hell (2012) Mitochondrial cysteine synthase complex regulates O-acetylserine biosynthesis in plants. Journal of Biological Chemistry 287, 27941-27947.

N.J. Kruger, S.K. Masakapalli and R.G. Ratcliffe (2012) Strategies for investigating the plant metabolic network with steady-state metabolic flux analysis: lessons from an Arabidopsis cell culture and other systems. Journal of Experimental Botany 63, 2309-2323.

N.J. Kruger and R.G. Ratcliffe (2012) Pathways and fluxes: exploring the plant metabolic network. Journal of Experimental Botany 63, 2243-2246.

T.C.R. Williams, L.J. Sweetlove and R.G. Ratcliffe (2011) Capturing metabolite channeling in metabolic flux phenotypes. Plant Physiology 157, 981-984.

L.J. Sweetlove and R.G. Ratcliffe (2011) Flux-balance modelling of plant metabolism. Frontiers in Plant Science 2:38. doi: 10.3389/fpls.2011.00038.

D.J.V. Beste, B. Bonde, N. Hawkins, J.L. Ward, M.H. Beale, S. Noack, K. Nöh, N.J. Kruger, R.G. Ratcliffe and J. McFadden (2011) 13C metabolic flux analysis identifies an unusual route for pyruvate dissimilation in mycobacteria which requires isocitrate lyase and carbon dioxide fixation. PLoS Pathogens 7, e1002091. doi:10.1371/journal.ppat.1002091

M. Watanabe, O. Yumi, Y. Itoh, K. Yasuda, K. Kamachi and R.G. Ratcliffe (2011) Deamination role of inducible glutamate dehydrogenase isoenzyme 7 in Brassica napus leaf protoplasts. Phytochemistry 72, 587-593.

Full Publication List (while at this department)

Kruger, N.J, Ratcliffe, R.G. (2015) Fluxes through plant metabolic networks: Measurements, predictions, insights and challenges Biochemical Journal. 465: pp 27-38.
doi:10.1042/BJ20140984.

Gupta, K.J, Mur, L.A.J, Ratcliffe, R.G. (2014) Guarding the guard cells? New Phytologist. 203 (2): pp 349-351.
doi:10.1111/nph.12882.

Igamberdiev, A.U, Ratcliffe, R.G, Gupta, K.J. (2014) Plant mitochondria: Source and target for nitric oxide Mitochondrion..
doi:10.1016/j.mito.2014.02.003.

Kruger, N.J, Masakapalli, S.K, Ratcliffe, R.G. (2014) Optimization of steady-state 13C-labeling experiments for metabolic flux analysis Methods in Molecular Biology. Eds. Dieuaide-Noubhani M.Alonso A.P.. ISBN: 9781627036870. pp 53-72.
doi:10.1007/978-1-62703-688-7-4

Masakapalli, S.K, Bryant, F.M, Kruger, N.J, Ratcliffe, R.G. (2014) The metabolic flux phenotype of heterotrophic Arabidopsis cells reveals a flexible balance between the cytosolic and plastidic contributions to carbohydrate oxidation in response to phosphate limitation Plant Journal. 78 (6): pp 964-977.
doi:10.1111/tpj.12522.

Masakapalli, S.K, Ritala, A, Dong, L, van, der Krol A.R, Oksman-Caldentey, K.-M, Ratcliffe, R.G, Sweetlove, L.J. (2014) Metabolic flux phenotype of tobacco hairy roots engineered for increased geraniol production Phytochemistry..
doi:10.1016/j.phytochem.2013.12.007

Masakapalli, S.K, Ratcliffe, R.G, Williams, T.C.R. (2014) Quantification of 13C enrichments and isotopomer abundances for metabolic flux analysis using 1D NMR spectroscopy Methods in Molecular Biology. Eds. Dieuaide-Noubhani M.Alonso A.P.. ISBN: 9781627036870. pp 73-86.
doi:10.1007/978-1-62703-688-7-5

Moller, B.L, Ratcliffe, R.G. (2014) Editorial overview: Synthetic plant biology: the roots of a bio-based society Current Opinion in Biotechnology..
doi:10.1016/j.copbio.2014.02.016

Cheung, C.Y.M, Williams, T.C.R, Poolman, M.G, Fell, D.A, Ratcliffe, R.G, Sweetlove, L.J. (2013) A method for accounting for maintenance costs in flux balance analysis improves the prediction of plant cell metabolic phenotypes under stress conditions Plant Journal..
doi:10.1111/tpj.12252

Masakapalli, S.K, Kruger, N.J, Ratcliffe, R.G. (2013) The metabolic flux phenotype of heterotrophic Arabidopsis cells reveals a complex response to changes in nitrogen supply Plant Journal..
doi:10.1111/tpj.12142

Sweetlove, L.J, Williams, T.C.R, Cheung, C.Y.M, Ratcliffe, R.G. (2013) Modelling metabolic CO2 evolution - a fresh perspective on respiration Plant, Cell and Environment..
doi:10.1111/pce.12105

Yang, Y.Y, Li, X.H, Ratcliffe, R.G, Ruan, J.Y. (2013) Characterization of ammonium and nitrate uptake and assimilation in roots of tea plants Russian Journal of Plant Physiology. 60 (1): pp 91-99.
doi:10.1134/S1021443712060180

Kruger, N.J, Ratcliffe, R.G. (2012) Pathways and fluxes: Exploring the plant metabolic network Journal of Experimental Botany. 63 (6): pp 2243-2246.
doi:10.1093/jxb/ers073.

Kruger, N.J, Masakapalli, S.K, Ratcliffe, R.G. (2012) Strategies for investigating the plant metabolic network with steady-state metabolic flux analysis: Lessons from an Arabidopsis cell culture and other systems Journal of Experimental Botany. 63 (6): pp 2309-2323.
doi:10.1093/jxb/err382

Wirtz, M, Beard, K.F.M, Lee, C.P, Boltz, A, Schwarzlander, M, Fuchs, C, Meyer, A.J, Heeg, C, Sweetlove, L.J, Ratcliffe, R.G, Hell, R. (2012) Mitochondrial cysteine synthase complex regulates O-acetylserine biosynthesis in plants Journal of Biological Chemistry. 287 (33): pp 27941-27947.
doi:10.1074/jbc.M112.372656

Beste, D.J.V, Bonde, B, Hawkins, N, Ward, J.L, Beale, M.H, Noack, S, Noh, K, Kruger, N.J, Ratcliffe, R.G, McFadden, J. (2011) 13c metabolic flux analysis identifies an unusual route for pyruvate dissimilation in mycobacteria which requires isocitrate lyase and carbon dioxide fixation PLoS Pathogens. 7 (7):.
doi:10.1371/journal.ppat.1002091

Watanabe, M, Yumi, O, Itoh, Y, Yasuda, K, Kamachi, K, Ratcliffe, R.G. (2011) Deamination role of inducible glutamate dehydrogenase isoenzyme 7 in Brassica napus leaf protoplasts Phytochemistry..
doi:10.1016/j.phytochem.2011.01.031

Masakapalli, S.K, le, Lay P, Huddleston, J.E, Pollock, N.L, Kruger, N.J, Ratcliffe, R George. (2010) Subcellular flux analysis of central metabolism in a heterotrophic Arabidopsis cell suspension using steady-state stable isotope labeling Plant Physiology. 152 (2): pp 602-619.
doi:10.1104/pp.109.151316

Sweetlove, L.J, Beard, K.F.M, Nunes-Nesi, A, Fernie, A.R, Ratcliffe, R.G. (2010) Not just a circle: flux modes in the plant TCA cycle Trends in Plant Science..
doi:10.1016/j.tplants.2010.05.006

Williams, T.C.R, Poolman, M.G, Howden, A.J.M, Schwarzlander, M, Fell, D.A, Ratcliffe, R.G, Sweetlove, L.J. (2010) A Genome-scale metabolic model accurately predicts fluxes in central carbon metabolism under stress conditions Plant Physiology. 154 (1): pp 311-323.
doi:10.1104/pp.110.158535

Allen, DK, Ratcliffe, RG. (2009) Quantification of Isotope Label. In Plant Metabolic Networks . Eds. J. Schwender. Springer, New York. ISBN: 978-0-387-78744-2 Print 978-0-387-78745-9 Online. pp 105-149.
doi:10.1007/978-0-387-78745-9_5

Couldwell, D.L, Dunford, R, Kruger, N.J, Lloyd, D.C, Ratcliffe, R.G, Smith, A.M.O. (2009) Response of cytoplasmic pH to anoxia in plant tissues with altered activities of fermentation enzymes: Application of methyl phosphonate as an NMR pH probe Annals of Botany. 103 (2): pp 249-258.
doi:10.1093/aob/mcn174

Kruger, N.J, Ratcliffe, R.G. (2009) Insights into plant metabolic networks from steady-state metabolic flux analysis Biochimie. 91 (6): pp 697-702.
doi:10.1016/j.biochi.2009.01.004

Troncoso-Ponce, M.A, Kruger, N.J, Ratcliffe, R.G, Garces, R, Martinez-Force, E. (2009) Characterization of glycolytic initial metabolites and enzyme activities in developing sunflower (Helianthus annuus L.) seeds. Phytochemistry. 70 (9): pp 1117-1122.
doi:10.1016/j.phytochem.2009.07.012

Kruger, N.J., Ratcliffe, R.G. (2008) Metabolic organization in plants: a challenge for the metabolic engineer. Advances in Plant Biochemistry and Molecular Biology : Bioengineering and Molecular Biology of Plant Pathways. 1: pp 1-27.
doi:10.1016/S1755-0408(07)01001-6

Kruger, N.J, Troncoso-Ponce, M.A, Ratcliffe, R.G. (2008) 1H NMR metabolite fingerprinting and metabolomic analysis of perchloric acid extracts from plant tissues Nature Protocols. 3 (6): pp 1001-1012.
doi:10.1038/nprot.2008.64

Morgan, M.J, Lehmann, M, Schwarzlander, M, Baxter, C.J, Sienkiewicz-Porzucek, A, Williams, T.C.R, Schauer, N, Fernie, A.R, Fricker, M.D, Ratcliffe, R.G, Sweetlove, L.J, Finkemeier, I. (2008) Decrease in manganese superoxide dismutase leads to reduced root growth and affects tricarboxylic acid cycle flux and mitochondrial redox homeostasis Plant Physiology. 147 (1): pp 101-114.
doi:10.1104/pp.107.113613

Williams, T.C.R, Miguet, L, Masakapalli, S.K, Kruger, N.J, Sweetlove, L.J, Ratcliffe, R.G. (2008) Metabolic network fluxes in heterotrophic arabidopsis cells: Stability of the flux distribution under different oxygenation conditions Plant Physiology. 148 (2): pp 704-718.
doi:10.1104/pp.108.125195

Graham, J.W.A, Williams, T.C.R, Morgan, M, Fernie, A.R, Ratcliffe, R.G, Sweetlove, L.J. (2007) Glycolytic enzymes associate dynamically with mitochondria in response to respiratory demand and support substrate channeling Plant Cell. 19 (11): pp 3723-3738.
doi:10.1105/tpc.107.053371

Kruger, N.J, Huddleston, J.E, Le, Lay P, Brown, N.D, Ratcliffe, R.G. (2007) Network flux analysis: Impact of 13C-substrates on metabolism in Arabidopsis thaliana cell suspension cultures Phytochemistry. 68: pp 2176-2188.
doi:10.1016/j.phytochem.2007.03.033

Kruger, N.J, Ratcliffe, R.G. (2007) Dynamic metabolic networks: Going with the flow Phytochemistry. 68: pp 2136-2138.
doi:10.1016/j.phytochem.2007.04.035

Kruger, N.J, Le, Lay P, Ratcliffe, R.G. (2007) Vacuolar compartmentation complicates the steady-state analysis of glucose metabolism and forces reappraisal of sucrose cycling in plants Phytochemistry. 68: pp 2189-2196.
doi:10.1016/j.phytochem.2007.04.004

Nunes-Nesi, A, Carrari, F, Gibon, Y, Sulpice, R, Lytovchenko, A, Fisahn, J, Graham, J, Ratcliffe, R.G, Sweetlove, L.J, Fernie, A.R. (2007) Deficiency of mitochondrial fumarase activity in tomato plants impairs photosynthesis via an effect on stomatal function Plant Journal. 50 (6): pp 1093-1106.
doi:10.1111/j.1365-313X.2007.03115.x

Dixon, M.H, Hill, S.A, Jackson, M.B, Ratcliffe, R.G, Sweetlove, L.J. (2006) Physiological and metabolic adaptations of Potamogeton pectinatus L. tubers support rapid elongation of stem tissue in the absence of oxygen Plant and Cell Physiology. 47 (1): pp 128-140.
doi:10.1093/pcp/pci229

Libourel, I.G.L, Van, Bodegom P.M, Fricker, M.D, Ratcliffe, R.G. (2006) Nitrite reduces cytoplasmic acidosis under anoxia Plant Physiology. 142 (4): pp 1710-1717.
doi:10.1104/pp.106.088898

Malone, J.G, Mittova, V, Ratcliffe, R.G, Kruger, N.J. (2006) The response of carbohydrate metabolism in potato tubers to low temperature Handbook of Environmental Chemistry, Volume 5: Water Pollution. 47 (9): pp 1309-1322.
doi:10.1093/pcp/pcj101

Ratcliffe, R.G, Shachar-Hill, Y. (2006) Measuring multiple fluxes through plant metabolic networks Plant Journal. 45 (4): pp 490-511.
doi:10.1111/j.1365-313X.2005.02649.x

Krishnan, P, Kruger, N.J, Ratcliffe, R.G. (2005) Metabolite fingerprinting and profiling in plants using NMR Journal of Experimental Botany. 56 (410): pp 255-265.
doi:10.1093/jxb/eri010

Mesnard, F, Ratcliffe, R.G. (2005) NMR analysis of plant nitrogen metabolism Photosynthesis Research. 83 (2): pp 163-180.
doi:10.1007/s11120-004-2081-8

Nunes-Nesi, A, Carrari, F, Lytovchenko, A, Smith, A.M.O, Loureiro, M.E, Ratcliffe, R.G, Sweetlove, L.J, Fernie, A.R. (2005) Enhanced photosynthetic performance and growth as a consequence of decreasing mitochondrial malate dehydrogenase activity in transgenic tomato plants Plant Physiology. 137 (2): pp 611-622.
doi:10.1104/pp.104.055566

Radcliffe, S.A, Miller, A.J, Ratcliffe, R.G. (2005) Microelectrode and 133Cs nuclear magnetic resonance evidence for variable cytosolic and cytoplasmic nitrate pools in maize root tips Plant, Cell and Environment. 28 (11): pp 1379-1387.
doi:10.1111/j.1365-3040.2005.01370.x

Ratcliffe, R.G, Shachar-Hill, Y. (2005) Revealing metabolic phenotypes in plants: Inputs from NMR analysis Biological Reviews of the Cambridge Philosophical Society. 80 (1): pp 27-43.
doi:10.1017/S1464793104006530

Smith, A.M.O, Ratcliffe, R.G, Sweetlove, L.J. (2004) Activation and function of mitochondrial uncoupling protein in plants Journal of Biological Chemistry. 279 (50): pp 51944-51952.
doi:10.1074/jbc.M408920200

Kruger, N.J, Ratcliffe, R.G, Roscher, A. (2003) Quantitative approaches for analysing fluxes through plant metabolic networks using NMR and stable isotope labelling Phytochemistry Reviews. 2: pp 17-30 

Stoimenova, M, Libourel, I.G.L, Ratcliffe, R.G, Kaiser, W.M. (2003) The role of nitrate reduction in the anoxic metabolism of roots II. Anoxic metabolism of tobacco roots with or without nitrate reductase activity Plant and Soil. 253 (1): pp 155-167.
doi:10.1023/A:1024591116697

Fernie, A.R, Roscher, A, Ratcliffe, R.G, Kruger, N.J. (2002) Activation of pyrophosphate:fructose-6-phosphate 1-phosphotransferase by fructose 2,6-bisphosphate stimulates conversion of hexose phosphates to triose phosphates but does not influence accumulation of carbohydrates in phosphate-deficient tobacco cells Physiologia Plantarum. 114 (2): pp 172-181.
doi:10.1034/j.1399-3054.2002.1140203.x

Gerendás, J, Ratcliffe, RG. (2002) Root pH regulation Plant Roots: The Hidden Half - 3rd edition, revised and expanded. pp 553-570 

Hartung, W, Leport, L, Ratcliffe, R.G, Sauter, A, Duda, R, Turner, N.C. (2002) Abscisic acid concentration, root pH and anatomy do not explain growth differences of chickpea (Cicer arietinum L.) and lupin (Lupinus angustifolius L.) on acid and alkaline soils Plant and Soil. 240 (1): pp 191-199.
doi:10.1023/A:1015831610452

Hartung, W, Ratcliffe, R.G. (2002) Utilization of glycine and serine as nitrogen sources in the roots of Zea mays and Chamaegigas intrepidus Journal of Experimental Botany. 53 (379): pp 2305-2314 

Mesnard, F, Roscher, A, Garlick, A.P, Girard, S, Baguet, E, Arroo, R.R.J, Lebreton, J, Robins, R.J, Ratcliffe, R.G. (2002) Evidence for the involvement of tetrahydrofolate in the demethylation of nicotine by Nicotiana plumbaginifolia cell-suspension cultures Planta. 214 (6): pp 911-919.
doi:10.1007/s00425-001-0702-4

Aubert, S, Bligny, R, Douce, R, Gout, E, Ratcliffe, R.G, Roberts, J.K.M. (2001) Contribution of glutamate dehydrogenase to mitochondrial glutamate metabolism studied by 13C and 31P nuclear magnetic resonance Journal of Experimental Botany. 52 (354): pp 37-45 

Dwek, RA, Wormald, M, Ratcliffe, RG, Price, NC. (2001) Principles and Problems in Physical Chemistry for Biochemists 3rd Edition . Oxford University Press, Oxford, UK. ISBN: 0-19-879281-6. 

Fernie, A.R, Roscher, A, Ratcliffe, R.G, Kruger, N.J. (2001) Fructose 2,6-bisphosphate activates pyrophosphate: Fructose-6-phosphate 1-phosphotransferase and increases triose phosphate to hexose phosphate cycling heterotrophic cells Planta. 212 (2): pp 250-263.
doi:10.1007/s004250000386

Ratcliffe, R.G, Roscher, A, Shachar-Hill, Y. (2001) Plant NMR spectroscopy Progress in Nuclear Magnetic Resonance Spectroscopy. 39 (4): pp 267-300.
doi:10.1016/S0079-6565(01)00035-8

Ratcliffe, R.G, Shachar-Hill, Y. (2001) Probing plant metabolism with NMR Annual Review of Plant Biology. 52: pp 499-526 

Ford, Y.-Y, Ratcliffe, R.G, Robins, R.J. (2000) An in vivo 15N NMR study of agropine synthesis in transformed root cultures of Nicotiana tabacum Physiologia Plantarum. 109 (2): pp 123-128.
doi:10.1034/j.1399-3054.2000.100203.x

Gerendas, J, Ratcliffe, R.G. (2000) Intracellular pH regulation in maize root tips exposed to ammonium at high external pH Journal of Experimental Botany. 51 (343): pp 207-219 

Heilmeier, H, Ratcliffe, RG, Hartung, W. (2000) Urea: A Nitrogen Source for the Aquatic Resurrection Plant Chamaegigas intrepidus Dinter Oecologia. 123 (1): pp 9-14 

Rasmussen, N, Lloyd, D.C, Ratcliffe, R.G, Hansen, P.E, Jakobsen, I. (2000) 31P NMR for the study of P metabolism and translocation in arbuscular mycorrhizal fungi Plant and Soil. 226 (2): pp 245-253.
doi:10.1023/A:1026411801081

Roscher, A, Kruger, N.J, Ratcliffe, R.G. (2000) Strategies for metabolic flux analysis in plants using isotope labelling Journal of Biotechnology. 77 (1): pp 81-102.
doi:10.1016/S0168-1656(99)00209-6

Summers, J.E, Ratcliffe, R.G, Jackson, M.B. (2000) Anoxia tolerance in the aquatic monocot Potamogeton pectinatus: Absence of oxygen stimulates elongation in association with an unusually large Pasteur effect Journal of Experimental Botany. 51 (349): pp 1413-1422 

Sweetlove, L.J, Dunford, R, Ratcliffe, R.G, Kruger, N.J. (2000) Lactate metabolism in potato tubers deficient in lactate dehydrogenase activity Plant, Cell and Environment. 23 (8): pp 873-881.
doi:10.1046/j.1365-3040.2000.00605.x

Kruger, NJ, Hill, SA, Ratcliffe, RG. (1999) Regulation of Primary Metabolic Pathways in Plants Proceedings of the Phytochemical Society of Europe (volume 42). Kluwer Academic Publishers, Dordrecht, The Netherlands. ISBN: 0-7923-5494-X. 

Ratcliffe, RG. (1999) Intracellular pH regulation in plants under anoxia Regulation of tissue pH in plants and animals: a reappraisal of current techniques. pp 193-213 

Ford, Y.-Y, Ratcliffe, R.G, Robins, R.J. (1998) In vivo nuclear-magnetic-resonance analysis of polyamine and alkaloid metabolism in transformed root cultures of Datura stramonium L.: Evidence for the involvement of putrescine in phytohormone-induced de-differentiation Planta. 205 (2): pp 205-213.
doi:10.1007/s004250050313

Ratcliffe, R.G, Roscher, A. (1998) Prospects for in vivo NMR methods in xenobiotic research in plants Biodegradation. 9 (6): pp 411-422.
doi:10.1023/A:1008372107739

Schiller, P, Hartung, W, Ratcliffe, R.G. (1998) Intracellular pH stability in the aquatic resurrection plant Chamaegigas intrepidus in the extreme environmental conditions that characterize its natural habitat New Phytologist. 140 (1): pp 1-7.
doi:10.1046/j.1469-8137.1998.00245.x

Gambhir, P.N, Pande, P.C, Ratcliffe, R.G. (1997) An in vivo 31P NMR study of the phosphorus metabolites in developing seeds of wheat, soybean and mustard Magnetic Resonance in Chemistry. 35: 

Gerendas, J, Zhu, Z, Bendixen, R, Ratcliffe, R.G, Sattelmacher, B. (1997) Physiological and biochemical processes related to ammonium toxicity in higher plants Journal of Plant Nutrition and Soil Science. 160 (3): pp 239-251 

Gerendas, J, Heeschen, V, Kahl, S, Ratcliffe, R.G, Rudolph, H. (1997) An investigation of N metabolism and pH regulation in Sphagnum using in vivo nuclear magnetic resonance and stable isotope mass spectrometry Isotopes in Environmental and Health Studies. 33 (1): pp 21-29 

Kahl, S, Gerendás, J, Heeschen, V, Ratcliffe, RG, Rudolph, H. (1997) Ammonium assimilation in bryophytes. L-glutamine synthetase from Sphagnum fallax Physiologia Plantarum. 101: pp 86-92 

Ratcliffe, R.G. (1997) In vivo nmr studies of the metabolic response of plant tissues to anoxia Annals of Botany. 79: pp 39-48 

Shachar-Hill, Y, Befroy, D.E, Pfeffer, P.E, Ratcliffe, R.G. (1997) Using Bulk Magnetic Susceptibility to Resolve Internal and External Signals in the NMR Spectra of Plant Tissues Journal of Magnetic Resonance. 127 (1): pp 17-25 

Ford, Y.-Y, Ratcliffe, R.G, Robins, R.J. (1996) Phytohormone-induced GABA production in transformed root cultures of Datura stramonium: An in vivo 15N NMR study Journal of Experimental Botany. 47 (299): pp 811-818 

Ford, Y.-Y, Ratcliffe, R.G, Robins, R.J. (1996) In vivo NMR analysis of tropane alkaloid metabolism in transformed root and de-differentiated cultures of Datura stramonium Phytochemistry. 43 (1): pp 115-120.
doi:10.1016/0031-9422(96)00261-0

Ratcliffe, RG. (1996) In vivo NMR spectroscopy: biochemical and physiological applications to plants Nuclear magnetic resonance in plant biology. pp 1-32 

Shachar-Hill, Y, Pfeffer, PE, Ratcliffe, RG. (1996) Measuring nitrate in plant cells by in vivo NMR using Gd3+ as a shift reagent Journal of Magnetic Resonance - Series B. 111 (1): pp 9-14 

Fox, G.G, McCallan, N.R, Ratcliffe, R.G. (1995) Manipulating cytoplasmic pH under anoxia: A critical test of the role of pH in the switch from aerobic to anaerobic metabolism Planta. 195 (3): pp 324-330 

Fox, G.G, Ratcliffe, R.G, Robinson, S.A, Stewart, G.R. (1995) Evidence for deamination by glutamate dehydrogenase in higher plants: Commentary Canadian Journal of Botany. 73 (7): pp 1112-1115 

Gerendás, J, Ratcliffe, RG, Sattelmacher, B. (1995) The influence of nitrogen and potassium supply on the ammonium content of maize (Zea mays L.) leaves including a comparison of measurements made in vivo and in vitro Plant and Soil. 173 (1): pp 11-20 

Quiquampoix, H, Abadie, J, Baron, MH, Leprince, F, Matumoto-Pintro, PT, Ratcliffe, RG, Staunton, S. (1995) Mechanisms and consequences of protein adsorption on soil mineral surfaces Proteins at interfaces II: fundamentals and applications (ACS Symposium Series). 602: pp 321-333 

Ratcliffe, RG. (1995) Metabolic aspects of the anoxic response in plant tissue Environment and plant metabolism: flexibility and acclimation. pp 111-127 

Shachar-Hill, Y, Pfeffer, PE, Douds, D, Osman, SF, Doner, LW, Ratcliffe, RG. (1995) Partitioning of carbohydrate metabolism in vesicular-arbuscular mycorrhizal leek Plant Physiology. 108 (1): pp 7-15 

Carroll, A.D, Fox, G.G, Laurie, S, Phillips, R, Ratcliffe, R.G, Stewart, G.R. (1994) Ammonium assimilation and the role of gamma-aminobutyric acid in pH homeostasis in carrot cell suspensions Plant Physiology. 106 (2): pp 513-520 

Ford, Y.Y, Fox, G.G, Ratcliffe, R.G, Robins, R.J. (1994) In vivo 15N NMR studies of secondary metabolism in transformed root cultures of Datura stramonium and Nicotiana tabacum Phytochemistry. 36 (2): pp 333-339.
doi:10.1016/S0031-9422(00)97071-7

Ratcliffe, R.G. (1994) In vivo NMR studies of higher plants and algae ADV.BOT.RES.. 20: pp 43-123 

Sklenar, J, Fox, G.G, Loughman, B.C, Pannifer, A.D.B, Ratcliffe, R.G. (1994) Effects of vanadate on the ATP content, ATPase activity and phosphate absorption capacity of maize roots Plant and Soil. 167 (1): pp 57-62.
doi:10.1007/BF01587598

Doyle, TC, Spickett, CM, Donaldson, IA, Ratcliffe, RG. (1993) Metabolic studies of a fructose-intolerant yeast by in vivo 31P nuclear magnetic resonance spectroscopy Yeast. 9 (8): pp 867-873 

Gerendas, J, Ratcliffe, R.G, Sattelmacher, B. (1993) Relationship between intracellular pH and N metabolism in maize (Zea mays L.) roots Plant and Soil. (1): pp 167-170.
doi:10.1007/BF00025010

Lee, R.B, Ratcliffe, R.G. (1993) Nuclear magnetic resonance studies of the location and function of plant nutrients in vivo Plant and Soil. (1): pp 45-55.
doi:10.1007/BF00024983

Lee, R.B, Ratcliffe, R.G. (1993) Observations on the subcellular distribution of the ammonium ion in maize root tissue using in-vivo 14N-nuclear-magnetic-resonance spectroscopy Planta. 189 (1): pp 156.
doi:10.1007/BF00201356

Quiquampoix, H, Staunton, S, Baron, M.-H, Ratcliffe, R.G. (1993) Interpretation of the pH dependence of protein adsorption on clay mineral surfaces and its relevance to the understanding of extracellular enzyme activity in soil Colloids and Surfaces A: Physicochemical and Engineering Aspects. 75: pp 85-93 

Quiquampoix, H, Loughman, B.C, Ratcliffe, R.G. (1993) A 31P-NMR study of the uptake and compartmentation of manganese by maize roots Journal of Experimental Botany. 44 (269): pp 1819-1827 

Quiquampoix, H, Bacic, G, Loughman, B.C, Ratcliffe, R.G. (1993) Quantitative aspects of the 31P-NMR detection of manganese in plant tissues Journal of Experimental Botany. 44 (269): pp 1809-1818 

Spickett, C.M, Smirnoff, N, Ratcliffe, R.G. (1993) An in vivo nuclear magnetic resonance investigation of ion transport in maize (Zea mays) and Spartina anglica roots during exposure to high salt concentrations Plant Physiology. 102 (2): pp 629-638 

Fox, G.G, Ratcliffe, R.G, Robinson, S.A, Slade, A.P, Stewart, G.R. (1992) Detection of 15N-labeled ammonium. In Vivo, 15N NMR versus mass spectrometry Journal of Magnetic Resonance (1969). 96 (1): pp 146-153 

Lee, RB, Purves, JV, Ratcliffe, RG, Saker, LR. (1992) Nitrogen assimilation and the control of ammonium and nitrate absorption by maize roots Journal of Experimental Botany. 43 (11): pp 1385-1396 

Obrenović, S, Ratcliffe, RG, Southon, TE. (1992) Direct observation of the effects of 3,5-diisopropylsalicylate and light on maize coleoptiles using 31P NMR Biochemie und Physiologie der Pflanzen. 188 (2): pp 136-143 

Quiquampoix, H, Ratcliffe, R.G. (1992) A 31P NMR study of the adsorption of bovine serum albumin on montmorillonite using phosphate and the paramagnetic cation Mn2+: modification of conformation with pH Journal of Colloid And Interface Science. 148 (2): pp 343-352 

Rands, ML, Douglas, AE, Loughman, BC, Ratcliffe, RG. (1992) Avoidance of hypoxia in a Cnidarian symbiosis by algal photosynthetic oxygen Biological Bulletin. 182: pp 159-162 

Spickett, C.M, Smirnoff, N, Ratcliffe, R.G. (1992) Metabolic response of maize roots to hyperosmotic shock: An in vivo31P nuclear magnetic resonance study Plant Physiology. 99 (3): pp 856-863 

Lee, R.B, Ratcliffe, R.G. (1991) Observations on the subcellular distribution of the ammonium ion in maize root tissue using in-vivo 14N-nuclear magnetic resonance spectroscopy Planta. 183 (3): pp 359-367.
doi:10.1007/BF00197734.

Ratcliffe, RG. (1991) Nuclear magnetic resonance in plant science research Botanical Journal of Scotland. 46: pp 107-120 

Robinson, S.A, Slade, A.P, Fox, G.G, Phillips, R, Ratcliffe, R.G, Stewart, G.R. (1991) The role of glutamate dehydrogenase in plant nitrogen metabolism Plant Physiology. 95 (2): pp 509-516 

Bright, AAS, Ratcliffe, RG. (1990) Application of NMR in the analysis of agrochemicals and pesticides Annual Reports on NMR Spectroscopy. 22: pp 139-203 

Fox, G.G, Ratcliffe, R.G. (1990) 31P NMR observations on the effect of the external pH on the intracellular pH values in plant cell suspension cultures Plant Physiology. 93 (2): pp 512-521 

Gerendás, J, Ratcliffe, RG, Sattelmacher, B. (1990) 31P nuclear magnetic resonance evidence for differences in intracellular pH in the roots of maize seedlings grown with nitrate or ammonium Journal of Plant Physiology. 137 (1): pp 125-128 

Lee, RB, Ratcliffe, RG, Southon, TE. (1990) 31P NMR measurements of the cytoplasmic and vacuolar Pi content of mature maize roots: relationships with phosphorus status and phosphate fluxes Journal of Experimental Botany. 41 (9): pp 1063-1078 

Quiquampoix, H, Ratcliffe, RG, Ratković, S, Vučinić, Z. (1990) A 1H and 31P NMR investigation of gadolinium uptake in maize roots Journal of Inorganic Biochemistry. 38 (4): pp 265-275 

Ratcliffe, RG, Roberts, JKM. (1990) Recent applications of NMR to higher plants and algae Magnetic Resonance in Medicine and Biology. 4 (1): pp 77-99 

Fox, G.G, Ratcliffe, R.G, Southon, T.E. (1989) Airlift systems for in vivo NMR spectroscopy of plant tissues Journal of Magnetic Resonance (1969). 82 (2): pp 360-366 

Fox, GG, Loughman, BC, Ratcliffe, RG, Rigden, DJ. (1989) Extending the NMR method to whole plants Plant Membrane Transport. pp 385-386 

Lines, CEM, Ratcliffe, RG, Rees, TAV, Southon, TE. (1989) A 13C NMR study of photosynthate transport and metabolism in the lichen Xanthoria calcicola Oxner New Phytologist. 111 (3): pp 447-456 

Loughman, B.C, Ratcliffe, R.G, Schwabe, J.W.R. (1989) Galactose metabolism in Zea mays root tissues observed by 31P-NMR spectroscopy Plant Science. 59 (1): pp 11-23 

Loughman, B.C, Ratcliffe, R.G, Southon, T.E. (1989) Observations on the cytoplasmic and vacuolar orthophosphate pools in leaf tissues using in vivo 31P-NMR spectroscopy FEBS Letters. 242 (2): pp 279-284 

Quiquampoix, H, Chassin, P, Ratcliffe, RG. (1989) Enzyme activity and cation exchange as tools for the study of the conformation of proteins adsorbed on mineral surfaces Progress in Colloid and Polymer Science. 79: pp 59-63 

Cairney, JWG, Jennings, DH, Ratcliffe, RG, Southon, TE. (1988) The physiology of basidiomycete linear organs. II. Phosphate uptake by rhizomorphs of Armillaria mellea New Phytologist. 109 (3): pp 327-333 

Ginzburg, M, Ratcliffe, R.G, Southon, T.E. (1988) Phosphorus metabolism and intracellular pH in the halotolerant alga Dunaliella parva studied by 31P-NMR BBA - Molecular Cell Research. 969 (3): pp 225-235 

Lohman, JAB, Ratcliffe, RG. (1988) Prospects for NMR imaging in the study of biological morphogenesis Experientia. 44 (8): pp 666-672

Connelly, A, Lohman, JAB, Loughman, BC, Quiquampoix, H, Ratcliffe, RG. (1987) High resolution imaging of plant tissues by NMR Journal of Experimental Botany. 38 (10): pp 1713-1723 

Ratcliffe, RG. (1987) The application of NMR methods to plant tissues Methods in Enzymology. 148: pp 683-700 

Ratcliffe, R.G. (1986) NMR and the inorganic composition of plants Journal of Inorganic Biochemistry. 28: pp 347-354 

Belton, PS, Ratcliffe, RG. (1985) NMR and compartmentation in biological tissues Progress in NMR Spectroscopy. 17 (2): pp 241-279 

Belton, PS, Lee, RB, Ratcliffe, RG. (1985) A 14N nuclear magnetic resonance study of inorganic nitrogen metabolism in barley, maize and pea roots Journal of Experimental Botany. 36 (163): pp 190-210 

Reid, RJ, Loughman, BC, Ratcliffe, RG. (1985) 31P NMR measurements of cytoplasmic pH changes in maize root tips Journal of Experimental Botany. 36 (6): pp 889-897

Thomas, TH, Ratcliffe, RG. (1985) 13C NMR determination of sugar levels in storage roots of carrot (Daucus carota, L.) Physiologia Plantarum. 63 (3): pp 284-286 

Loughman, BC, Ratcliffe, RG. (1984) Nuclear magnetic resonance and the study of plants Advances in Plant Nutrition. 1: pp 241-283 

Robins, RJ, Ratcliffe, RG. (1984) Intracellular distribution of phosphate in cultured Humulus lupulus cells growing at elevated exogenous phosphate concentrations Plant Cell Reports. 3 (6): pp 234-236