The DOI matches its Crossref record; 59 of the 64 references it cites were found in a registry or an index.
A Universal Definition of Life: Autonomy and Open-Ended Evolution
Abstract
Life is a complex phenomenon that not only requires individual self-producing and self-sustaining systems but also a historical-collective organization of those individual systems, which brings about characteristic evolutionary dynamics. On these lines, we propose to define universally living beings as autonomous systems with open-ended evolution capacities, and we claim that all such systems must have a semi-permeable active boundary (membrane), an energy transduction apparatus (set of energy currencies) and, at least, two types of functionally interdependent macromolecular components (catalysts and records). The latter is required to articulate a 'phenotype-genotype' decoupling that leads to a scenario where the global network of autonomous systems allows for an open-ended increase in the complexity of the individual agents. Thus, the basic-individual organization of biological systems depends critically on being instructed by patterns (informational records) whose generation and reliable transmission cannot be explained but take into account the complete historical network of relationships among those systems. We conclude that a proper definition of life should consider both levels, individual and collective: living systems cannot be fully constituted without being part of the evolutionary process of a whole ecosystem. Finally, we also discuss a few practical implications of the definition for different programs of research.
Overview
Integrity signals
Registry
Cites
Statistics
Verification
Claims the paper makes
Citations
Cites
The Origins of Order: Self-Organization and Selection in Evolution
Kauffman, S. A. (1993) · doi:10.1093/oso/9780195079517.001.0001 · Cited by 7,855
The origins of order; self organization and selection in evolution
Kauffman, S. A. (1994) · International Journal of Biochemistry · doi:10.1016/0020-711x(94)90119-8 · Cited by 6,753
Theory of Self-Reproducing Automata
Schwartz, J. T., von Neumann, J., & Burks, A. W. (1967) · Mathematics of Computation · doi:10.2307/2005041 · Cited by 5,468
THE MAJOR TRANSITIONS OF EVOLUTION
Leigh, E. G., Smith, J. M., & Szathmáry, E. (1995) · Evolution · doi:10.1111/j.1558-5646.1995.tb04464.x · Cited by 3,435
The Major Transitions in Evolution
Smith, J. M., & Szathmáry, E. (1997) · Oxford University Press eBooks · doi:10.1093/oso/9780198502944.001.0001 · Cited by 2,570
Lewontin, R. C. (1970) · Annual Review of Ecology and Systematics · doi:10.1146/annurev.es.01.110170.000245 · Cited by 1,996
The Hypercycle: A principle of natural self-organization
Eigen, M., & Schuster, P. M. (1979) · Cited by 1,906
Phylogenetic Classification and the Universal Tree
Doolittle, W. F. (1999) · Science · doi:10.1126/science.284.5423.2124 · Cited by 1,888
Principles of Biological Autonomy
Varela, F. J. (2025) · The MIT Press eBooks · doi:10.7551/mitpress/14875.001.0001 · Cited by 1,876
Szostak, J. W., Bartel, D. P., & Luisi, P. L. (2001) · Nature · doi:10.1038/35053176 · Cited by 1,681
Atmospheric homeostasis by and for the biosphere: the gaia hypothesis
Lovelock, J. E., & Margulis, L. (1974) · Tellus · doi:10.3402/tellusa.v26i1-2.9731 · Cited by 1,386
Before enzymes and templates: theory of surface metabolism
Wächtershäuser, G. (1988) · Microbiological Reviews · doi:10.1128/mmbr.52.4.452-484.1988 · Cited by 1,242
Life itself: a comprehensive inquiry into the nature, origin, and fabrication of life
(1992) · Choice Reviews Online · doi:10.5860/choice.29-4505 · Cited by 1,043
Kauffman, S. A. (2000) · doi:10.1093/oso/9780195121049.001.0001 · Cited by 950
Eigen, M., & Schuster, P. (1979) · doi:10.1007/978-3-642-67247-7 · Cited by 949
Dyson, F. J. (1985) · Cited by 615
Acquiring Genomes: A Theory Of The Origins Of Species
Margulis, L., & Sagan, D. (2002) · Cited by 424
Compositional genomes: Prebiotic information transfer in mutually catalytic noncovalent assemblies
Segré, D., Ben-Eli, D., & Lancet, D. (2000) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.97.8.4112 · Cited by 398
Healey, I. N., & Morowitz, H. J. (1970) · Journal of Ecology · doi:10.2307/2258194 · Cited by 368
Cleland, C. E., & Chyba, C. F. (2002) · Origins of life and evolution of the biosphere · doi:10.1023/a:1020503324273 · Cited by 347
Evolution as Entropy: Toward a Unified Theory of Biology
Geesink, R., Brooks, D. R., & WILEY, E. O. (1986) · Taxon · doi:10.2307/1221696 · Cited by 325
Koshland, D. E. (2002) · Science · doi:10.1126/science.1068489 · Cited by 314
Dynamic and linguistic modes of complex systems
PATTEE, H. H. (1977) · International Journal of General Systems · doi:10.1080/03081077708934771 · Cited by 273
Blueprint for a cell - the nature and origin of life
(1991) · Choice Reviews Online · doi:10.5860/choice.28-5078 · Cited by 273
About Various Definitions of Life
Luisi, P. L. (1998) · Origins of life and evolution of the biosphere · doi:10.1023/a:1006517315105 · Cited by 266
Code-duality and the semiotics of nature
Hoffmeyer, J., & Emmeche, C. (1991) · De Gruyter eBooks · doi:10.1515/9783110849875.117 · Cited by 265
The chemical logic of a minimum protocell
Morowitz, H. J., Heinz, B., & Deamer, D. W. (1988) · Origins of life and evolution of the biosphere · doi:10.1007/bf01804674 · Cited by 239
Life : its nature, origin and development
Oparin, A. I. (1962) · Academic Press eBooks · Cited by 196
Evolution, Thermodynamics, and Information: Extending the Darwinian Program
Wicken, J. S. (1987) · Medical Entomology and Zoology · Cited by 190
Smith, J. M. (1986) · Medical Entomology and Zoology · Cited by 189
Life detection by atmospheric analysis
Hitchcock, D. R., & Lovelock, J. E. (1967) · Icarus · doi:10.1016/0019-1035(67)90059-0 · Cited by 166
Steps towards life: a perspective on evolution
(1993) · Choice Reviews Online · doi:10.5860/choice.30-3812 · Cited by 164
Evolution as Entropy. Toward a Unified Theory of Biology
Fauchald, K., Brooks, D. R., & WILEY, E. O. (1987) · Journal of Parasitology · doi:10.2307/3282343 · Cited by 142
John von Neumann and the Evolutionary Growth of Complexity: Looking Backward, Looking Forward
McMullin, B. (2000) · Artificial Life · doi:10.1162/106454600300103674 · Cited by 124
Evolution, thermodynamics and information
Wicken, J. S. (1987) · Cited by 121
De máquinas y seres vivos: Una teoría sobre la organización biológica
Maturana, H. R., & Varela, F. J. (1973) · Cited by 116
Pályi, G., Zucchi, C., & Caglioti, L. (2002) · Cited by 92
Artificial life: The coming evolution
Farmer, J. D., Belin, A. D., & Farmer, J. D. (1990) · Cited by 84
Metabolism and the problem of its universalization
Bergareche, A. M., & Ruiz-Mirazo, K. (1999) · Biosystems · doi:10.1016/s0303-2647(98)00034-3 · Cited by 78
Self-citationAutopoiesis: the status of its system logic
Fleischaker, G. R. (1988) · Biosystems · doi:10.1016/0303-2647(88)90048-2 · Cited by 75
Defining life as a semiotic phenomenon.
Emmeche, C. (1998) · Research at the University of Copenhagen (University of Copenhagen) · Cited by 63
Quantum mechanics, heredity and the origin of life
Pattee, H. H. (1967) · Journal of Theoretical Biology · doi:10.1016/0022-5193(67)90101-4 · Cited by 51
Membrane compartments in prebiotic evolution
Deamer, D. W. (1998) · Cambridge University Press eBooks · doi:10.1017/cbo9780511626180.010 · Cited by 50
Homeostatic Tendencies of the Earth’s Atmosphere
Lovelock, J. E., & Margulis, L. (1974) · Springer eBooks · doi:10.1007/978-94-010-2239-2_8 · Cited by 47
Searching for the roots of autonomy: The natural and artificial paradigms revisited
Ruiz‐Mirazo, K., & Moreno, Á. (2000) · Cited by 41
Self-citationPRICE, M. L. (1905) · JAMA · doi:10.1001/jama.1905.52510110015001b · Cited by 36
Semantic Closure: A guiding notion to ground artificial life
Umerez, J. (1995) · Lecture notes in computer science · doi:10.1007/3-540-59496-5_290 · Cited by 31
The Chemical Implementation of Autopoiesis
Luisi, P. L. (1994) · Springer eBooks · doi:10.1007/978-94-011-0754-9_15 · Cited by 30
Phase separation, charge separation and biogenesis
Morowitz, H. J. (1981) · Biosystems · doi:10.1016/0303-2647(81)90020-4 · Cited by 20
Bro, P. (1997) · Complexity · doi:10.1002/(sici)1099-0526(199701/02)2:3 · Cited by 20
Possible Forms of Life in Environments Very Different from the Earth
Shapiro, R., & Feinberg, G. (1995) · Cambridge University Press eBooks · doi:10.1017/cbo9780511564970.020 · Cited by 17
Possible forms of life in environments very different from the Earth.
Shapiro, R., & Feinberg, G. (1982) · Cited by 13
Lazcano, A. (2001) · Wiley eBooks · doi:10.1002/9780470999295.ch1 · Cited by 13
Varela, F. J. (1994) · Springer eBooks · doi:10.1007/978-94-011-0754-9_2 · Cited by 12
Moreno, Á., & Ruiz‐Mirazo, K. (2002) · Emergence · doi:10.1080/15213250.2002.9687735 · Cited by 10
Self-citationA Few Precautionary Words Concerning Terminology
Fleischaker, G. R. (1994) · Springer eBooks · doi:10.1007/978-94-011-0754-9_3 · Cited by 9
Life as emergence: The roots of a new paradigm in theoretical biology
Fernandez, J., Moreno, Á., & Etxeberría, A. (1991) · World Futures · doi:10.1080/02604027.1991.9972255 · Cited by 9
Sagan, D., & Margulis, L. (1997) · Springer eBooks · doi:10.1007/978-1-4612-2284-2_11 · Cited by 8
Condiciones físicas para la aparición de sistemas autónomos con capacidades evolutivas abiertas
Mirazo, P. R. (2001) · Euskomedia Digital Bilduma · Cited by 4
Research works printed but not linked: 15
- Benner, S. A.: 1999, How Small can a Microorganism be? in Size Limits of Very Small Microorganisms (Proceedings of a Workshop), National Academy Press, Washington, D.C., pp. 126–135.
- Haynes, R. H.: 1990, ‘Ecce Ecopoiesis: Playing God on Mars’, in D. MacNiven (ed.), Moral Expertise: Studies in Practical and Professional Ethics, Routledge, London, pp. 161–183.
- Joyce, G. F.: 1994, ‘Foreword’, in D. W. Deamer and G. R. Fleischaker (eds), Origins of Life: The Central Concepts, Jones and Bartlett, Boston, pp. xi–xii.
- Lovelock, J. E.: 1988, The Ages of Gaia, Norton, New York.
- Margulis, L.: 1990, ‘Big Trouble in Biology: Physiological Autopoiesis Versus Mechanistic Neo-Darwinism’, in J. Brockman (ed.), Doing Science: The Reality Club 2, Prentice Hall, NY, pp. 211–235.
- Mayr, E.: 1982, The Growth of Biological Thought, Harvard University Press, Cambridge, MA.
- Moreno, A. and Fernández, J.: 1990, Structural Limits for Evolutive Capacities in Molecular Complex Systems, Biology Forum 83, 335–347.
- Moreno, A., Umerez, J. and Fernandez, J.: 1994, Definition of Life and Research Program in Artificial Life, Ludus Vitalis 2, 15–33.
- Morowitz, H. J.: 1992, Beginnings of Cellular Life, Yale University Press, New Haven.
- Nicolis, G. and Prigogine, Y.: 1977, Self-organization in Non-equilibrium Systems, Wiley, New York.
- Pattee, H. H.: 1982, Cell Psychology: An Evolutionary Approach to the Symbol-matter Problem, Cognit. Brain Theor. 4, 325–341.
- Pattee, H. H.: 1997, ‘The Physics of Symbols and Evolution of Semiotic Controls’, in M. Coombs (ed.), Workshop on Control Mechanisms for Complex Systems: Issues of Measurement and Semiotic Analysis, New Mexico State University.
- Ruiz-Mirazo, K. and Moreno, A.: 1998, ‘Autonomy and Emergence: how Systems become Agents through the Generation of Functional Constraints, in G. L. Farre and T. Oksala (eds), Emergence, Complexity, Hierarchy, Organization (Selected and edited papers from the ECHO III Conference), Acta Polytech. Scand. Ma91, The Finnish Academy of Technology, Espoo-Helsinki, pp. 273–282.
- Sagan, C.: 1970, ‘Life’, in The Encyclopaedia Britannica, William Benton, London.
- Shapiro, R.: 2002, ‘Monomer World’, in Abstracts of the 13th International Conference on the Origin of Life, 10th ISSOL Meeting, Oaxaca, Mexico, p. 60.
Cited by
Sorted by citations
Explanation: a mechanist alternative
Bechtel, W., & Abrahamsen, A. A. (2005) · Studies in History and Philosophy of Science Part C Studies in History and Philosophy of Biological and Biomedical Sciences · doi:10.1016/j.shpsc.2005.03.010 · Cited by 1,143
Prebiotic Systems Chemistry: New Perspectives for the Origins of Life
Ruiz-Mirazo, K., Briones, C., & de la Escosura, A. (2014) · Chemical Reviews · doi:10.1021/cr2004844 · Cited by 930
Self-citationLearning how to live together: genomic insights into prokaryote–animal symbioses
Moyá, A., Peretó, J. G., Gil, R., & Latorre, A. (2008) · Nature Reviews Genetics · doi:10.1038/nrg2319 · Cited by 544
Self-citationDefining Agency: Individuality, Normativity, Asymmetry, and Spatio-temporality in Action
Barandiaran, X. E., Di Paolo, E., & Rohde, M. (2009) · Adaptive Behavior · doi:10.1177/1059712309343819 · Cited by 497
Origins of building blocks of life: A review
Kitadai, N., & Maruyama, S. (2018) · Geoscience Frontiers · doi:10.1016/j.gsf.2017.07.007 · Cited by 464
Metadata
- Authors
- Ruiz-Mirazo, K., Peretó, J., & Moreno, A.
- Affiliations
- Universitat de València (ES)
- ORCID
- Peretó, Juli
- Venue
- Origins of life and evolution of the biosphere
- Published
- 2004-06
- Volume
- 34
- Issue
- 3
- Pages
- 323-346
- ISSN
- 0169-6149 (ISSN-L)
- 1573-0875
- OpenAlex
- W2089114542
- Cited by
- 393
- References
- 64
- Citations by year
- 2026: 17 · 2025: 16 · 2024: 22 · 2023: 17 · 2022: 13 · 2021: 32 · 2020: 13 · 2019: 40 · 2018: 19 · 2017: 18 · 2016: 15 · 2015: 13 · 2014: 31 · 2013: 21 · 2012: 13
- FWCI
- 7.28
- Type
- journal article
- Language
- en
- Open access
- closed
- Licence
- Locations
- Origins of Life and Evolution of Biospheres (publishedVersion)
- PubMed (publishedVersion)
- Funders
- Comisión Interministerial de Ciencia y Tecnología
- Instituto Nacional de Técnica Aeroespacial
- Topics
- Physical Sciences › Physics and Astronomy › Astronomy and Astrophysics › Origins and Evolution of Life
- Social Sciences › Arts and Humanities › History and Philosophy of Science › Philosophy of Biology and Scientific Explanation
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Evolution and Genetic Dynamics
- Keywords
- concept of life, autonomy, open-ended evolution, semipermeable membranes, energy transduction, collective organization, evolutionary dynamics, complexity increase
- MeSH terms
- Origin of Life, Energy Metabolism, Biological Evolution, Models, Biological, Models, Theoretical, Reproduction, Life
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review