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Theoretical Biology of Systems

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This is the home page for the research group of Athel Cornish-Bowden and María Luz Cárdenas at the CNRS in Marseilles, now part of the group on Extremophile Metabolism directed by Marie-Thérèse Giudici-Orticoni.

The 4th edition of Athel Cornish-Bowden (2012) Fundamentals of Enzyme Kinetics was published by Wiley–Blackwell on 25th January 2012. Details of the changes in the new edition may be found on another page.

A new and extensively revised edition of Athel Cornish-Bowden (1995) The Pursuit of Perfection was published in May 2016, by Garland Science, New York & London, under a new title: Biochemical Evolution: the Pursuit of Perfection. More information can be found here.

 Cover design      Cover design


Members of the group

 [Photograph] Marilú Athel

María Luz Cárdenas and Athel Cornish-Bowden

Research interests

Our recent publications are listed below. Complete publication lists are also available, arranged either by year of publication or under various broad headings, as follows:

  1. Current or recent activities
  2. Earlier research interests
  3. Books from our group

A list of our most highly cited publications may be found on another page.


Bioénergétique et Ingénierie des Protéines, Centre National de la Recherche Scientifique, 31 chemin Joseph-Aiguier, B.P. 71, 13402 Marseille Cedex 20, France

 [Picture of the Vieux Port]


The laboratory Bioénergétique et Ingénierie des Protéines (BIP, CNRS UMR 7281) is a Unité Mixte de Recherche of the CNRS and Aix-Marseille Université. It is directed by Dr Marie-Thérèse Giudici-Orticoni and is a component unit of the Groupe de Laboratoires de Marseille du CNRS (Marseilles Group of CNRS Laboratories).

Getting here

A separate web page gives suggestions on how to reach the CNRS.

Contact information

Current publications

Complete lists may be found here (AJCB) or here (MLC). This list includes only articles in press or recently published. Links to the corresponding entries in the full lists are given in square brackets. There is also a (very long) complete list, organized according to date of publication.

PDF files of most papers published in journals are directly linked from this page, as indicated with the symbol [ Download PDF ]. Some other PDF files (marked with the symbol [ Request PDF ]) are available on request to If a [ DOI link ] symbol is given it is a link to the publisher’s site.


A. Cornish-Bowden & M. L. Cárdenas (2020) Contrasting Theories of Life: Historical Context, Current Thories. In Search of an Ideal Theory BioSystems 188, 104063 [ DOI link ] 10.1016/j.biosystems.2019.104063 Until 11 March 2020 a PDF file can be downloaded without cost and without registering at If this doesn't work (as it may not if your browser cannot handle https links or has other problems) please contact me at


C. Mariscal, A. Barahona, N. Aubert-Kato, A. U. Aydinoglu, S. Bartlett, M. L. Cárdenas, K. Chandru, C. Cleland, B. T. Cocanougher, N. Comfort, A. Cornish-Bowden, T. Deacon, T. Froese, D. Giovannelli, J. Hernlund, P. Hut, J. Kimura, M.-C. Maurel, N. Merino, A. Moreno, M. Nakagawa, J. Peretó, N. Virgo, O. Witkowski, & H. J. Cleaves II Hidden Concepts in the History and Philosophy of Origins-of-Life Studies: A Workshop Report Orig. Life Evol. Biosph. 49, 111 [ DOI link ] 10.1007/s11084-019-09580-x


M. L. Cárdenas, S. Benomar & A. Cornish-Bowden (2018) Rosennean complexity and its relevance to ecology Ecological Complexity 35, 13–24 [Mulitienzyme systems (MLC), Mu48 and Life (AJCB), J23] [ Download PDF ] [ DOI link ] 10.1016/j.ecocom.2017.04.005

N. Swainston, A. Baici, B. M. Bakker, A. Cornish-Bowden, P. F. Fitzpatrick, P. Halling, T. S. Leyh, C. O'Donovan, F. M. Raushel, J. M. Rohwer, S. Schnell, D. Schomburg, K. F. Tipton, M.-D. Tsai, H. V. Westerhoff, U. Wittig, R. Wohlgemuth & C. Kettner (2018) STRENDA DB: enabling the validation and sharing of enzyme kinetics data: enabling the validation and sharing of enzyme kinetics data FEBS J. 285, 2193–2214 [ Download PDF ] [ DOI link ] 10.1111/febs.14427


A. Cornish-Bowden Overview of the 3rd Conference on Aneuploidy and Cancer in Berkeley, CA USA in 2017 Mol. Cytogen.10 (supplement 2), 77-80 [ Download PDF ] [ DOI link ] 10.1186/s13039-017-0320-x

A. Cornish-Bowden Enthalpy–entropy compensation and the isokinetic temperature in enzyme catalysis J. Biosci. 42, pp. 665–670 [ DOI link ] 10.1007/s12038-017-9719-0

A. Cornish-Bowden & M. L. Cárdenas (2017) Life before LUCA J. Theor. Biol. 434, 68–74 [Mulitienzyme systems (MLC), Mu49 and Life (AJCB), J23] [ Download PDF ] [ DOI link ] 10.1016/j.jtbi.2017.05.023


A. Bocedi, R. Fabrini, M. Lo Bello, A. M. Caccuri, G. Federici, B. Mannervik, A. Cornish-Bowden & G. Ricci (2016) Evolution of negative cooperativity in glutathione transferase enabled preservation of enzyme function J. Biol. Chem. 292, 26739–26749 [Other enzymes, F9. See also Multienzyme systems, Hf8 and Ligand binding, B7] [ Download PDF ] [ DOI link ] 10.1074/jbc.M116.749507

M. L. Cárdenas (2016) Profesor Giorgio Semenza (1928–2016) SEBBM No. 188, 88–49 [ Download PDF ] [Miscellaneous, Mm18]

A. Cornish-Bowden (2016) Biochemical Evolution: the Pursuit of Perfection Garland Science (263 pp.) New York & London [See also the first edition]


S. Benomar, D. Ranava, M. L. Cárdenas, E. Trably, Y. Rafrafi, A. Ducret, J. Hamelin, E. Lojou, J.-P. Steyer & M. T. Giudici-Orticoni (2015) Nutritional stress induces exchange of cell material and energetic coupling between bacterial species Nat. Comm. 6, 6283 [ Download PDF ] [ DOI link ] 10.1038/ncomms7283 Supplementary material [Miscellaneous, Mm17]

M. L. Cárdenas (2015) Understanding mechanisms of enzyme cooperativity: the importance of not being at equilibrium Persp. Sci. 4, 10–16 [ Download PDF ] [ DOI link ] 10.1016/j.pisc.2014.12.003 [Enzyme kinetics, Mk12]

A. Cornish-Bowden (2015) One hundred years of Michaelis–Menten kinetics Persp. Sci. 4, 3–9 [ Download PDF ] [ DOI link ] 10.1016/j.pisc.2014.12.002

A. Cornish-Bowden (2015) Tibor Gánti and Robert Rosen: contrasting approaches to the same problem J. Theor. Biol. 381, 6–10 [ Request PDF ] [ DOI link ] 10.1016/j.jtbi.2015.05.015 [Life, J22]

A. Cornish-Bowden (2015) Борис Гольдштейн и его вывд уравнений скорости стацонарных реакций [ Download PDF ] (in Russian: translated from "Boris Goldstein and the Derivation of Steady-State Rate Equations" [ Download PDF ] ) pp. 125–138 in Папа: Пущино. Профессия. Палиндромы (ed. L. Lavrova) Nauchniy Mir, Moscow [Kinetic analysis, C57]


J. L. Asenjo, H. C. Ludwig, C. A. Droppelmann, J. G. Cárcamo, I. I. Concha, A. J. Yáñez, M. L. Cárdenas, A. Cornish-Bowden & J. C. Slebe (2014) Subunit interactions in pig-kidney fructose-1,6-bisphosphatase: binding of substrate induces a second class of site with lowered affinity and catalytic activity Biochim. Biophys. Acta 1840, 1798–1807 [ Download PDF ] [ DOI link ] 10.1016/j.bbagen.2013.12.027 [Enzyme kinetics, Mk11 and Other enzymes, F8]

A. Cornish-Bowden (2014) Understanding allosteric and cooperative interactions in enzymes FEBS J. 281, 621–632 [ Download PDF ] [ DOI link ] 10.1111/febs.12469 [Ligand binding, B11]

A. Cornish-Bowden (2014) Professor Robert Arnold Alberty (1921–2014) The Biochemist 36(3), 57 [ Download PDF ] [Miscellaneous, K35]

A. Cornish-Bowden (2014) Current IUBMB recommendations on enzyme nomenclature and kinetics Persp. Sci. 1, 74–87 [ Download PDF ] [ DOI link ] 10.1016/j.pisc.2014.02.006 [Kinetic analysis, C53]

A. Cornish-Bowden (2014) Analysis and interpretation of enzyme kinetic data Persp. Sci. 1, 121–125 [ Download PDF ] [ DOI link ] 10.1016/j.pisc.2014.02.010 [Kinetic analysis, C54]

A. Cornish-Bowden & C. Kettner (2014) Persp. Sci. 1, 1–137 (organizers of a special issue entitled Reporting Enzymology Data — STRENDA Recommendations and Beyond). All of the papers are available at from

A. Cornish-Bowden, J.-P. Mazat & S. Nicolas (2014) Victor Henri: 111 years of his equation Biochimie 107, 161–166 [ Download PDF ] Supplementary material: Thesis in English; Thesis in French (six parts), Prelims, Introduction, Chapter 1, Chapter 2, Chapter 3, Chapter 4 to end. [ DOI link ] 10.1016/j.biochi.2014.09.018 [Kinetic analysis, C56]

A. Cornish-Bowden, J. Peretó & M. L. Cárdenas (2014) Biochemistry and evolutionary biology: two disciplines that need each other J. Biosci. 39, 1–15 [ Download PDF ] [ DOI link ] 10.1007/s12038-014-9414-3 [Miscellaneous (AJCB), K34 and Miscellaneous (MLC), Mm16]

G. Curien, M. L. Cárdenas & A. Cornish-Bowden (2014) Analytical kinetic modelling – a practical procedure in Methods in Molecular Biology (ed. M. Dieuaide-Noubhani & A. P. Alonso) 1090, 261–280 [Simulation, I13 and Multienzyme systems, Mu47]

U. Deichmann, S. Schuster, J.-P. Mazat & A. Cornish-Bowden (2014) Commemorating the 1913 Michaelis–Menten paper Die Kinetik der Invertinwirkung: three perspectives FEBS J. 281, 435–463 [ Download PDF ] [ DOI link ] 10.1111/febs.12598 [Miscellaneous, K33]

C. Kettner & A. Cornish-Bowden (2014) Quo Vadis, enzymology data? Introductory remarks (Introduction to Reporting Enzymology Data — STRENDA Recommendations and Beyond), Persp. Sci. 1, 1–6 [ Download PDF ] [ DOI link ] 10.1016/j.pisc.2014.02.013 [ Kinetic analysis, C52]

K. F. Tipton, R. N. Armstrong, B. M. Bakker, A. Bairoch, A. Cornish-Bowden, P. J. Halling, J.-H. Hofmeyr, T. S. Leyh, C. Kettner, F. M. Raushel, J. Rohwer, D. Schomburg & C. Steinbeck (2014) Standards for reporting enzyme data: the STRENDA consortium: what it aims to do and why it should be helpful Persp. Sci. 1, 131–137 [ Download PDF ] [ DOI link ] 10.1016/j.pisc.2014.02.012 [ Kinetic analysis, C46] [Life, J19]

Earlier years

Books from our Group

Athel Cornish-Bowden (1995) Analysis of Enzyme Kinetic Data, Oxford University Press, Oxford. [ Download PDF ]

María Luz Cárdenas (1995) "Glucokinase": its Regulation and Role in Liver Metabolism, R. G. Landes, Austin, Texas

Athel Cornish-Bowden, ed. (1997) New Beer in an Old Bottle: Eduard Buchner and the Growth of Biochemical Knowledge, Universitat de València, Valencia

Athel Cornish-Bowden (1999) Basic Mathematics for Biochemists (2nd edn.), Oxford University Press, Oxford

Athel Cornish-Bowden & María Luz Cárdenas (2000) Technological and Medical Implications of Metabolic Control Analysis, Kluwer Academic Publishers, Dordrecht, The Netherlands

Athel Cornish-Bowden (2004) The Pursuit of Perfection Oxford University Press, Oxford

Athel Cornish-Bowden, Marc Jamin & Valdur Saks (2005) Cinétique Enzymatique (in French), EDP Sciences, Paris

Athel Cornish-Bowden (2012) Fundamentals of Enzyme Kinetics (4th edn.), Wiley-VCH, Weinheim

Athel Cornish-Bowden (2016) Biochemical Evolution: the Pursuit of Perfection, Garland Science, New York

Erdős numbers

Each of us has an Erdős number of 3, based on the following chain:

A. Cornish-Bowden, M. L. Cárdenas, J.-C. Letelier, J. Soto-Andrade and F. Guíñez Abarzúa (2004) "Understanding the parts in terms of the whole" Biology of the Cell 96, 713–717

Wen-Ch'ing Winnie Li and J. Soto-Andrade (1983) "Barnes' identities and representations of GL (2). I. Finite field case" Journal für die reine und angewandte Mathematik 344, 171–179

S. A. Burr, P. Erdos, R. L. Graham and W. Li (1996) "Complete sequences of sets of integer powers" Acta Arithmetica 77, 133–138

Current or recent research collaborations

Metabolic control

General information on metabolic control analysis.

Interview on Beilstein TV. If that gives an error message (some servers don't allow local mp4 files), try the Beilstein site.