Skip to main content Accessibility help
×
Hostname: page-component-7dd5485656-jtdwj Total loading time: 0 Render date: 2025-10-31T05:22:44.075Z Has data issue: false hasContentIssue false

What Is Life? Revisited

Published online by Cambridge University Press:  10 October 2025

Daniel J. Nicholson
Affiliation:
George Mason University

Summary

Erwin Schrödinger's What Is Life? is one of the most celebrated scientific works of the twentieth century. However, like most classics, it is far more often cited than read. Efforts to seriously engage with Schrödinger's arguments are rare. This Element explores how well his ideas have stood the test of time. It argues that Schrödinger's emphasis on the rigidity and specificity of the hereditary material (which stemmed from his attempt to explain biological order from physical principles) influenced how molecular biologists conceptualized macromolecules, resulting in a deterministic, engineering view of the cell that is still popular today—even if it is increasingly at odds with experimental findings. Drawing on archival sources, this Element also uncovers Schrödinger's motivations in writing What Is Life? and suggests that his biological proposals are best understood in the context of his longstanding dispute with other physicists regarding the interpretation and extension of quantum mechanics.
Get access

Information

Type
Element
Information
Online ISBN: 9781009127318
Publisher: Cambridge University Press
Print publication: 13 November 2025

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

Element purchase

Temporarily unavailable

References

Abir-Am, P. 1985. Themes, genres, and orders of legitimation in the consolidation of new scientific disciplines: Deconstructing the historiography of molecular biology. History of Science 23: 73–117.CrossRefGoogle ScholarPubMed
Akam, M. 1989. Making stripes inelegantly. Nature 341: 282–283.CrossRefGoogle ScholarPubMed
Alberts, B. 1998. The cell as a collection of protein machines: Preparing the next generation of molecular biologists. Cell 92: 291–294.CrossRefGoogle ScholarPubMed
Alexander, J. and Bridges, C. B. 1929. Some physicochemical aspects of life, mutation and evolution. Science 70: 508–510.CrossRefGoogle ScholarPubMed
Anon. 1938. Prof. E. Schrödinger and the University of Graz. Nature 141: 929.Google Scholar
Ball, P. 2011. The dawn of quantum biology. Nature 474: 272–274.CrossRefGoogle ScholarPubMed
Ball, P. 2015. Forging patterns and making waves from biology to geology: A commentary on Turing (1952) ‘The chemical basis of morphogenesis’. Philosophical Transactions of the Royal Society B 370: 20140218. http://dx.doi.org/10.1098/rstb.2014.0218CrossRefGoogle ScholarPubMed
Ball, P. 2018. In retrospect: What Is Life? Nature 560: 548–550.Google Scholar
Ball, P. 2023. How Life Works: A User’s Guide to the New Biology. University of Chicago Press.CrossRefGoogle Scholar
Bębenek, A, and Ziuzia-Graczyk, I. 2018. Fidelity of DNA replication – A matter of processing. Current Genetics 64: 985–996.CrossRefGoogle Scholar
Benzer, S. 2002. Interview by Heidi Aspaturian. Oral History Project, California Institute of Technology Archives. https://digital.archives.caltech.edu/collections/OralHistories/OH_Benzer_S/Google Scholar
Bernstein, J. 2016. A Bouquet of Numbers and Other Scientific Offerings. World Scientific.CrossRefGoogle Scholar
Beyler, R. H. 1994. From positivism to organicism: Pascual Jordan’s interpretations of modern physics in cultural context. Dissertation, Harvard University.Google Scholar
Beyler, R. H. 1996. Targeting the organism: The scientific and cultural context of Pascual Jordan’s quantum biology, 1932–1947. Isis 87: 248–273.Google Scholar
Bialek, W., Cavagna, A., Giardina, I., Mora, T., Edmondo, S., Viale, M., and Walczak, A. M. 2012. Statistical mechanics for natural flocks of birds. Proceedings of the National Academy of Sciences 109: 4786–4791.CrossRefGoogle ScholarPubMed
Bohr, N. 1999. Collected Works, Vol. 10: Complementarity Beyond Physics (1928–1962). Elsevier.Google Scholar
Boltzmann, L. 1974. Theoretical Physics and Philosophical Problems: Selected Writings. Reidel.CrossRefGoogle Scholar
Bonduriansky, R. and Day, T. 2018. Extended Heredity: A New Understanding of Inheritance and Evolution. Princeton University Press.Google Scholar
Boycott, A. E. 1929. The transition from live to dead: The nature of filtrable viruses. Nature 123: 91–98.CrossRefGoogle Scholar
Brabazon. 1945. Causality or indeterminism? Nature 155: 398.Google Scholar
Brioullin, L. 1949. Life, thermodynamics, and cybernetics. American Scientist 37: 554–568.Google Scholar
Brioullin, L. 1956. Science and Information Theory. Academic Press.CrossRefGoogle Scholar
Butler, J. A. V. 1946. Life and the second law of thermodynamics. Nature 158: 153–154.CrossRefGoogle ScholarPubMed
Carlson, E. A. 1971. An unacknowledged founding of molecular biology: H. J. Muller’s contributions to gene theory, 1910–1936. Journal of the History of Biology 4: 149–170.CrossRefGoogle Scholar
Ceccarelli, L. 2001. Shaping Science with Rhetoric: The Cases of Dobzhansky, Schrödinger, and Wilson. University of Chicago Press.CrossRefGoogle Scholar
Cairns, J., Stent, G. S., and Watson, J. D. 1966. Phage and the Origins of Molecular Biology. Cold Spring Harbor Laboratory Press.Google Scholar
Chargaff, E. 1978. Heraclitean Fire: Sketches from a Life Before Nature. Rockefeller University Press.Google Scholar
Clary, D. C. 2022. Schrödinger in Oxford. World Scientific.CrossRefGoogle Scholar
Cobb, M. 2015. Life’s Greatest Secret: The Race to Crack the Genetic Code. Profile.Google Scholar
Crick, F. H. C. 1958. On protein synthesis. Symposium of the Society for Experimental Biology 12: 138–163.Google ScholarPubMed
Crick, F. H. C. 1965. Recent research in molecular biology: Introduction. British Medical Bulletin 21: 183–186.CrossRefGoogle Scholar
Crick, F. H. C. 1966. Of Molecules and Men. University of Washington Press.Google Scholar
Crow, J. F. 1992. Erwin Schrödinger and the hornless cattle problem. Genetics 130: 237–239.CrossRefGoogle ScholarPubMed
Darlington, C. D. and Mather, K. 1950. Genes, Plants and People: Essays on Genetics. Allen & Unwin.Google Scholar
Davidson, E. H. 2009. Q&A. Current Biology 22: R216–R217.Google Scholar
Davies, P. 2019. The Demon in the Machine: How Hidden Webs of Information Are Solving the Mystery of Life. University of Chicago Press.CrossRefGoogle Scholar
Dawkins, R. 1976. The Selfish Gene. Oxford University Press.Google Scholar
de Chadarevian, S. 1998. Of worms and programmes: Caenorhabditis elegans and the study of development. Studies in History and Philosophy of Biological and Biomedical Sciences 29: 81–105.CrossRefGoogle Scholar
Deacon, T. W. 2011. Incomplete Nature: How Mind Emerged from Matter. W. W. Norton.Google Scholar
Delbrück, M. 1945. What Is Life? and what is truth? Quarterly Review of Biology 20: 370–372.CrossRefGoogle Scholar
Delbrück, M. 1949. A physicist looks at biology. Transactions of the Connecticut Academy of Arts and Sciences 38: 173–190.Google Scholar
Depew, D. J. and Weber, B. H. 1995. Darwinism Evolving: Systems Dynamics and the Geneaology of Natural Selection. MIT Press.Google Scholar
Descartes, R. 1998. The World and Other Writings. Cambridge University Press.CrossRefGoogle Scholar
Donnan, F. G. 1918. La science physico-chimique décrit-elle d’une façon adéquate les phénomènes biologiques? Scientia 24: 282–288.Google Scholar
Donnan, F. G. 1928. The mystery of life. Journal of Chemical Education 122: 1558–1570.Google Scholar
Donnan, F. G. 1936. Integral analysis and the phenomena of life. Acta Biotheoretica 2: 1–11.CrossRefGoogle Scholar
Dowdle, E. B. 1989. Physics, Schrödinger and the study of life. Transactions of the Royal Society of South Africa 47: 103–108.CrossRefGoogle Scholar
Dronamraju, K. R. 1999. Erwin Schrödinger and the origins of molecular biology. Genetics 153: 1071–1076.CrossRefGoogle ScholarPubMed
Dyson, F. 1985. Origins of Life. Cambridge University Press.Google Scholar
Eldar, A. and Elowitz, M. B. 2010. Functional roles for noise in genetic circuits. Nature 467: 167–173.CrossRefGoogle ScholarPubMed
Elitzur, A. C. 1995. Life and mind, past and future: Schrödinger’s vision fifty years later. Perspectives in Biology and Medicine 38: 433–458.CrossRefGoogle Scholar
Fischer, E. P. 1984. We are all aspects of one single being: An introduction to Erwin Schrödinger. Social Research 51: 809–835.Google ScholarPubMed
Fisher, R. A. 1930. The Genetical Theory of Natural Selection. Oxford University Press.CrossRefGoogle Scholar
Fleming, D. 1968. Émigré physicists and the biological revolution. Perspectives in American History 2: 152–189.Google Scholar
Frank, J. 2011. Molecular Machines in Biology: Workshop of the Cell. Cambridge University Press.CrossRefGoogle Scholar
Fuller, S. 2021. Schrödinger’s What Is Life? as postdigital prophecy. Postdigital Science and Education 3: 272–279.CrossRefGoogle Scholar
Gamow, G. 1953. Mr. Tompkins Learns the Facts of Life. Cambridge University Press.CrossRefGoogle Scholar
Gamow, G. 1955. Information transfer in the living cell. Scientific American 193: 70–78.CrossRefGoogle Scholar
García-Ojalvo, J. and Martínez Arias, A. 2012. Towards a statistical mechanics of cell fate decisions. Current Opinion in Genetics & Development 22: 619–626.CrossRefGoogle ScholarPubMed
Gaskell, A. 1928. What Is Life? Thomas Books.Google Scholar
Gehring, W. J. 1988. Master Control Genes in Development and Evolution: The Homeobox Story. Yale University Press.Google Scholar
Gierer, A. and Meinhardt, H. 1972. A theory of biological pattern formation. Kybernetik 12: 30–39.CrossRefGoogle ScholarPubMed
Gilbert, W. 1992. A vision of the grail. In Kevles, D. J. and Hood, L. (eds.), The Code of Codes: Scientific and Social Issues in the Human Genome Project. Harvard University Press, pp. 83–97.Google Scholar
Gnaiger, E., Gellerich, F. N., and Wyss, M. 1994. What Is Controlling Life? 50 Years After Erwin Schrödinger’s What Is Life? Innsbruck University Press.Google Scholar
Gould, S. J. 1995. What Is Life? as a problem in history. In Murphy, M. P. and O’Neill, L. A. J. (eds.), What Is Life? The Next Fifty Years. Cambridge University Press, pp. 25–39.Google Scholar
Gribbin, J. 2013. Erwin Schrödinger and the Quantum Revolution. Wiley.Google Scholar
Green, J. B. A. and Sharpe, J. 2015. Positional information and reaction-diffusion: Two big ideas in developmental biology combine. Development 142: 1203–1211.CrossRefGoogle ScholarPubMed
Hacking, I. 1990. The Taming of Chance. Cambridge University Press.CrossRefGoogle Scholar
Haldane, J. B. S. 1945. A physicist looks at genetics. Nature 155: 375–376.CrossRefGoogle Scholar
Halpern, P. 2015. Einstein’s Dice and Schrödinger’s Cat: How Two Great Minds Battled Quantum Randomness to Create a Unified Theory of Physics. Basic Books.Google Scholar
Harrison, L. G. 1987. What is the status of reaction-diffusion theory thirty-four years after Turing? Journal of Theoretical Biology 125: 369–384.CrossRefGoogle Scholar
Heams, T. 2014. Randomness in biology. Mathematical Structures in Computer Science 24: 1–24.CrossRefGoogle Scholar
Heitler, W. 1961. Erwin Schrödinger (1887–1961). Biographical Memoirs of Fellows of the Royal Society 7: 221–228.Google Scholar
Hendrickson, M. R. 2011. Exorcizing Schrödinger’s ghost: Reflections on What Is Life? and its surprising relevance to cancer biology. In Gumbrecht, H. U., Harrison, R. P., Hendrickson, M. R., and Laughlin, R. B. (eds.), What Is Life? The Intellectual Pertinence of Erwin Schrödinger. Stanford University Press, pp. 45–103.Google Scholar
Hetler, D. M. and Bronfenbrenner, J. 1929. On the particulate size of bacteriophage. Proceedings of the Society for Experimental Biology and Medicine 26: 644–645.Google Scholar
Hoch, P. K. and Yoxen, E. J. 1987. Schrödinger at Oxford: A hypothetical national cultural synthesis which failed. Annals of Science 44: 593–616.CrossRefGoogle Scholar
Hodge, M. J. S. 1992. Biology and philosophy (including ideology): A study of Fisher and Wright. In Sarkar, S. (ed.), The Founders of Evolutionary Genetics. Kluwer, pp. 231–293.Google Scholar
Huang, S. 2009. Non-genetic heterogeneity of cells in development: More than just noise. Development 136: 3853–3862.CrossRefGoogle ScholarPubMed
Jablonka, E. and Lamb, M. J. 2014. Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life (rev. ed.). MIT Press.CrossRefGoogle Scholar
Jacob, F. 1973. The Logic of Life: A History of Heredity. Pantheon.Google Scholar
Jacob, F. 1988. The Statue Within: An Autobiography. Basic Books.Google Scholar
Jacob, F. and Monod, J. 1961. Genetic regulatory mechanisms in the synthesis of proteins. Journal of Molecular Biology 3: 318–356.CrossRefGoogle ScholarPubMed
Johannsen, W. 1923. Some remarks about units in heredity. Hereditas 4: 133–141.Google Scholar
Jordan, P. 1932. Die Quantenmechanik und die Grundprobleme der Biologie und Psychologie. Die Naturwissenschaften 20: 815–821.CrossRefGoogle Scholar
Judson, H. F. 1979. The Eighth Day of Creation: Makers of the Revolution in Biology. Simon & Schuster.Google Scholar
Kalmus, H. 1950. A cybernetical aspect of genetics. Journal of Heredity 41: 19–22.CrossRefGoogle Scholar
Karsenti, E. 2008. Self-organization in cell biology: A brief history. Nature Reviews Molecular Cell Biology 9: 255–262.CrossRefGoogle ScholarPubMed
Kauffman, S. 1995. What Is Life?: Was Schrödinger right? In Murphy, M. P. and O’Neill, L. A. J. (eds.), What Is Life? The Next Fifty Years. Cambridge University Press, pp. 83–114.Google Scholar
Kauffman, S. 2000. Investigations. Oxford University Press.CrossRefGoogle Scholar
Kay, L. E. 1985. The secret of life: Niels Bohr’s influence on the biology program of Max Delbrück. Rivista di Storia della Scienza 2: 487–510.Google Scholar
Kay, L. E. 1993. The Molecular Vision of Life: Caltech, the Rockefeller Foundation, and the Rise of the New Biology. Oxford University Press.Google Scholar
Kay, L. E. 2000. Who Wrote the Book of Life? A History of the Genetic Code. Stanford University Press.CrossRefGoogle Scholar
Keller, E. F. 1990. Physics and the emergence of molecular biology: A history of cognitive and political synergy. Journal of the History of Biology 23: 389–409.CrossRefGoogle ScholarPubMed
Keller, E. F. 1995. Refiguring Life: Metaphors of Twentieth-Century Biology. Columbia University Press.CrossRefGoogle Scholar
Keller, E. F. 2000. The Century of the Gene. Harvard University Press.CrossRefGoogle Scholar
Keller, E. F. 2002. Making Sense of Life: Explaining Biological Development with Models, Metaphors, and Machines. Harvard University Press.CrossRefGoogle Scholar
Keller, E. F. 2008. Organisms, machines, and thunderstorms: A history of self-organization, part one. Historical Studies in the Natural Sciences 38: 45–75.CrossRefGoogle Scholar
Keller, E. F. 2009. Organisms, machines, and thunderstorms: A history of self-organization, part two. Historical Studies in the Natural Sciences 39: 1–31.CrossRefGoogle Scholar
Kendrew, J. C. 1967. How molecular biology started. Scientific American 216: 141–144.Google Scholar
Kilmister, C. W. 1987. Schrödinger: Centenary Celebration of a Polymath. Cambridge University Press.CrossRefGoogle Scholar
Kirschner, M., Gerhart, M., and Mitchison, T. 2000. Molecular ‘vitalism’. Cell 100: 79–88.CrossRefGoogle ScholarPubMed
Kogge, W. 2012. Script, code, information: How to differentiate analogies in the ‘prehistory’ of molecular biology. History and Philosophy of the Life Sciences 34: 604–635.Google ScholarPubMed
Kondo, S. and Miura, T. 2010. Reaction-diffusion model as a framework for understanding biological pattern formation. Science 329: 1616–1620.CrossRefGoogle ScholarPubMed
Kupiec, J.-J. 2009. The Origin of Individuals. World Scientific.CrossRefGoogle Scholar
Kupiec, J.-J. 2010. On the lack of specificity of proteins and its consequences for a theory of biological organization. Progress in Biophysics and Molecular Biology 102: 45–52.CrossRefGoogle ScholarPubMed
Laughlin, R. B. 2011. Schrödinger’s trouble: How quantum mechanics got created with a logical loose end. In Gumbrecht, H. U., Harrison, R. P., Hendrickson, M. R., and Laughlin, R. B. (eds.), What Is Life ? The Intellectual Pertinence of Erwin Schrödinger. Stanford University Press, pp. 33–43.Google Scholar
Lewontin, R. C. 1968. Essay review: Phage and the origins of molecular biology. Journal of the History of Biology 1: 155–161.Google Scholar
Lewontin, R. C. 1970. The units of selection. Annual Review of Ecology and Systematics 1: 1–18.CrossRefGoogle Scholar
Lewontin, R. C. 2000. The Triple Helix: Gene, Organism, and Environment. Harvard University Press.Google Scholar
Loison, L. 2015. Why did Jacques Monod make the choice of mechanistic determinism? Comptes Rendus Biologies 338: 391–397.CrossRefGoogle Scholar
Longo, G. and Tendero, P. E 2007. The differential method and the causal incompleteness of programming theory in molecular biology. Foundations of Science 12: 337–366.CrossRefGoogle Scholar
Lovelock, J. A. 1986. Living alternatives. Nature 320: 646.CrossRefGoogle Scholar
Lovelock, J. A. 1988. The Ages of Gaia: A Biography of Our Living Earth. Oxford University Press.Google Scholar
Lwoff, A. 1962. Biological Order. MIT Press.CrossRefGoogle Scholar
MacArthur, B. D. and Lemischka, I. R. 2013. Statistical mechanics of pluripotency. Cell 154: 484–489.CrossRefGoogle ScholarPubMed
MacKenzie, D. A. 1981. Statistics in Britain (1865–1930): The Social Construction of Scientific Knowledge. Edinburgh University Press.Google Scholar
Maini, P. K. 2004. The impact of Turing’s work on pattern formation in biology. Mathematics Today 40: 140–141.Google Scholar
Manton, I. 1945. Comments on chromosome structure. Nature 155: 471–473.CrossRefGoogle Scholar
Margulis, L. and Sagan, D. 1995. What Is Life? University of California Press.Google Scholar
Maynard Smith, J. 1968. Mathematical Ideas in Biology. Cambridge University Press.CrossRefGoogle Scholar
Mayr, E. 1961. Cause and effect in biology. Science 134: 1501–1506.CrossRefGoogle ScholarPubMed
Mayr, E. 1997. This Is Biology: The Science of the Living World. Harvard University Press.Google Scholar
McFadden, J. and Al-Khalili, J. 2014. Life on the Edge: The Coming of Age of Quantum Biology. Crown Publishers.Google Scholar
McKaughan, D. J. 2005. The influence of Niels Bohr on Max Delbrück: Revisiting the hopes inspired by “Light and Life”. Isis 96: 507–529.CrossRefGoogle Scholar
Medawar, P. 1965. A biological retrospect. Nature 207: 1327–1330.CrossRefGoogle ScholarPubMed
Mertens, R. 2019. The Construction of Analogy-Based Research Programs: The Lock-and-Key Analogy in 20th Century Biochemistry. Transcript Verlag.Google Scholar
Moberg, C. 2020. Schrödinger’s What Is Life? – The 75th anniversary of a book that inspired biology. Angewandte Chemie 59: 2550–2553.Google ScholarPubMed
Monod, J. 1966. L’être vivant comme machine. In Caillois, R. (ed.), Le Robot, la Bête et l’Homme. Baconnière, pp. 43–47.Google Scholar
Monod, J. 1972. Chance and Necessity: An Essay on the Natural Philosophy of Molecular Biology. New Vintage.Google Scholar
Monod, J. 2021 [1960]. Cybérnetique Enzymatique. Èditions Matériologiques.Google Scholar
Moore, W. 1989. Schrödinger: Life and Thought. Cambridge University Press.CrossRefGoogle Scholar
Moore, P. B. 2012. How should we think about the ribosome? Annual Review of Biophysics 41: 1–19.CrossRefGoogle ScholarPubMed
Morange, M. 2008. Life Explained. Odile Jacob.Google Scholar
Morange, M. 2020. The Black Box of Biology: A History of the Molecular Revolution. Harvard University Press.CrossRefGoogle Scholar
Morowitz, H. J. 1970. Entropy for Biologists. Academic Press.Google Scholar
Moss, L. 2003. What Genes Can’t Do. MIT Press.Google Scholar
Muller, H. J. 1922. Variation due to change in the individual gene. American Naturalist 56: 32–50.CrossRefGoogle Scholar
Muller, H. J. 1929. The gene as the basis of life. Proceedings of the International Congress of Plant Sciences 1: 897–921.Google Scholar
Muller, H. J. 1937. Physics in the attack on the fundamental problems of genetics. Scientific Monthly 44: 210–214.Google Scholar
Muller, H. J. 1946. A physicist stands amazed at genetics. Journal of Heredity 32: 90–92.Google Scholar
Murphy, M. P. and O’Neill, L. A. J. 1995. What Is Life? The Next Fifty Years. Cambridge University Press.CrossRefGoogle Scholar
Needham, J. 1936. Order and Life. Cambridge University Press.Google Scholar
Newman, S. A. 1988. Idealist biology. Perspectives in Biology and Medicine 31: 353–368.CrossRefGoogle ScholarPubMed
Nicholson, D. J. 2014. The machine conception of the organism in development and evolution: A critical analysis. Studies in History and Philosophy of Biological and Biomedical Sciences 48: 162–174.CrossRefGoogle ScholarPubMed
Nicholson, D. J. 2018. Reconceptualizing the organism: From complex machine to flowing stream. In Nicholson, D. J. and Dupré, J. (eds.), Everything Flows: Towards a Processual Philosophy of Biology. Oxford University Press, pp. 139–166.CrossRefGoogle Scholar
Nicholson, D. J. 2019. Is the cell really a machine? Journal of Theoretical Biology 477: 108–126.CrossRefGoogle Scholar
Nicholson, D. J. 2020. On being the right size, revisited: The problem with engineering metaphors in molecular biology. In Holm, S. and Serban, M. (eds.), Philosophical Perspectives on the Engineering Approach in Biology: Living Machines? Routledge, pp. 40–68.Google Scholar
Nicolis, G. and Prigogine, I. 1977. Self-Organization in Non-Equilibrium Systems. Wiley.Google Scholar
Nijhout, H. F. 1990. Metaphors and the role of genes in development. BioEssays 12: 441–446.CrossRefGoogle ScholarPubMed
Noble, D. 2016. Dance to the Tune of Life – Biological Relativity. Cambridge University Press.Google Scholar
Nurse, P. 2021. What Is Life? Five Great Ideas in Biology. W. W. Norton.Google Scholar
O’Dwyer, J. P. 2020. Beyond an ecological ideal gas law. Nature Ecology & Evolution 4: 14–15.Google ScholarPubMed
Olby, R. 1970. Francis Crick, DNA, and the Central Dogma. Daedalus 99: 938–987.Google Scholar
Olby, R. 1971. Schrödinger’s problem: What is life? Journal of the History of Biology 4: 119–148.CrossRefGoogle ScholarPubMed
Olby, R. 1974. The Path to the Double Helix. University of Washington Press.Google Scholar
Oparin, A. I. 1961. Life: Its Nature, Origin and Development. Oliver & Boyd.Google Scholar
Oyama, S., Griffiths, P. E., and Gray, R. D. 2001. Cycles of Contingency: Developmental Systems and Evolution. MIT Press.Google Scholar
Pauling, L. 1987. Schrödinger’s contribution to chemistry and biology. In Kilmister, C. W. (ed.), Schrödinger: Centenary Celebration of a Polymath. Cambridge University Press, pp. 225–233.Google Scholar
Pence, C. H. 2022. The Rise of Chance in Evolutionary Theory: A Pompous Parade of Arithmetic. Academic Press.Google Scholar
Perutz, M. F. 1987a. Physics and the riddle of life. Nature 326: 555–558.CrossRefGoogle ScholarPubMed
Perutz, M. F. 1987b. Erwin Schrödinger’s What Is Life? and molecular biology. In Kilmister, C. W. (ed.), Schrödinger: Centenary Celebration of a Polymath. Cambridge University Press, pp. 234–251.Google Scholar
Phillips, R. 2021. Schrödinger’s What Is Life? at 75. Cell Systems 12: 465–476.CrossRefGoogle ScholarPubMed
Pollard, E. C. 1967. Erwin Schrödinger (1887–1961). In Snell, F. M. (ed.), Progress in Theoretical Biology, Vol. 1. Academic Press, pp. ix–xi.Google Scholar
Porter, T. M. 1986. The Rise of Statistical Thinking (1820–1900). Princeton University Press.CrossRefGoogle Scholar
Prigogine, I. and Stengers, I. 1984. Order Out of Chaos: Man’s New Dialogue with Nature. Bantam.Google Scholar
Pross, A. 2012. What Is Life? How Chemistry Becomes Biology. Oxford University Press.Google Scholar
Quastler, H. 1953. Essays on the Use of Information Theory in Biology. University of Illinois Press.Google Scholar
Regis, E. 2009. What Is Life? Investigating the Nature of Life in the Age of Synthetic Biology. Oxford University Press.CrossRefGoogle Scholar
Reitz, J. R. and Longmire, C. 1950. Living matter and physical laws. Physics Today 3: 15–19.CrossRefGoogle Scholar
Roll-Hansen, N. 2000. The application of complementarity to biology: From Niels Bohr to Max Delbrück. Historical Studies in the Physical and Biological Sciences 30: 417–442.CrossRefGoogle Scholar
Rose, S. 1998. Lifelines: Biology Beyond Determinism. Oxford University Press.Google Scholar
Rosen, R. 1996. The Schrödinger question: What is life? Fifty years later. In Buckley, P. and Peat, F. D. (eds.). Glimpsing Reality: Ideas in Physics and the Link to Biology. University of Toronto Press, pp. 168–190.Google Scholar
Rosenberg, A. 1997. Reductionism redux: Computing the embryo. Biology and Philosophy 12: 445–470.CrossRefGoogle Scholar
Sarkar, S. 1991. What Is Life? revisited. BioScience 41: 631–634.CrossRefGoogle Scholar
Sarkar, S. 1996. Biological information: A skeptical look at some central dogmas of molecular biology. In Sarkar, S. (ed.), The Philosophy and History of Molecular Biology. Springer, pp. 187–231.Google Scholar
Sarkar, S. 2013. Erwin Schrödinger’s excursus on genetics. In Harman, O. and Dietrich, M. R. (eds.), Outsider Scientists: Routes to Innovation in Biology. University of Chicago Press, pp. 93–109.Google Scholar
Schneider, E. D. 1987. Schrödinger’s grand theme shortchanged. Nature 328: 300.CrossRefGoogle ScholarPubMed
Schneider, E. D. and Kay, J. J. 1994. Life as a manifestation of the second law of thermodynamics. Mathematical and Computer Modeling 19: 25–48.CrossRefGoogle Scholar
Schneider, E. D. and Sagan, D. 2005. Into the Cool: Energy Flow, Thermodynamics, and Life. University of Chicago Press.Google Scholar
Schoenheimer, R. 1942. The Dynamic State of Body Constituents. Harvard University Press.Google Scholar
Schrödinger, E. 1933. Warum sind die Atome so klein? Forschungen und Fortschritte 9: 125–126.Google Scholar
Schrödinger, E. 1935a. Die gegenwärtige Situation in der Quantenmechanik. Naturwissenschaften 23: 807–812, 823–828, 844–849.CrossRefGoogle Scholar
Schrödinger, E. 1935b. Science and the Human Temperament. Allen & Unwin.Google Scholar
Schrödinger, E. 1936. Indeterminism and free will. Nature 138: 13–14.CrossRefGoogle Scholar
Schrödinger, E. 1944. What Is Life? The Physical Aspect of the Living Cell. Cambridge University Press.Google Scholar
Schrödinger, E. 1946. Statistical Thermodynamics. Cambridge University Press.Google Scholar
Schrödinger, E. 1951. Science and Humanism: Physics in Our Time. Cambridge University Press.Google Scholar
Schrödinger, E. 1954. Nature and the Greeks. Cambridge University Press.Google Scholar
Schrödinger, E. 1958. Mind and Matter. Cambridge University Press.Google Scholar
Schrödinger, E. 1964. My View of the World. Cambridge University Press.Google Scholar
Schrödinger, E. 1992. What Is Life? with Mind and Matter and Autobiographical Sketches. Cambridge University Press.CrossRefGoogle Scholar
Semon, R. 1904. Die Mneme als erhaltendes Prinzip. Engemann.Google Scholar
Shannon, C. 1956. The bandwagon. IRE Transactions on Information Theory IT 2: 3.Google Scholar
Shannon, C. and Weaver, W. 1949. The Mathematical Theory of Communication. University of Illinois Press.Google Scholar
Shostak, S. 1998. Death of Life: The Legacy of Molecular Biology. Macmillan.CrossRefGoogle Scholar
Sigmund, K. 2019. The physicist and the dawn of the double helix. Science 366: 43.Google Scholar
Sloan, P. R. 2012. How was teleology eliminated in early molecular biology? Studies in History and Philosophy of Biological and Biomedical Sciences 43: 140–151.CrossRefGoogle ScholarPubMed
Sloan, P. R. 2025. A return to Niels Bohr’s ‘Light and Life’ (1932). Biological Theory. https://doi.org/10.1007/s13752-025-00490-yCrossRefGoogle Scholar
Sloan, P. R. and Fogel, B. 2011. Creating a Physical Biology: The Three-Man Paper and Early Molecular Biology. University of Chicago Press.CrossRefGoogle Scholar
Stent, G. S. 1966. Introduction: Waiting for the paradox. In Cairns, J., Stent, G. S., and Watson, J. D. (eds.), Phage and the Origins of Molecular Biology. Cold Spring Harbor Laboratory Press, pp. 3–8.Google Scholar
Stent, G. S. 1968. That was the molecular biology that was. Science 160: 390–395.CrossRefGoogle ScholarPubMed
Sulston, J. E., Schierenberg, E., White, J. G., and Thomson, J. N. 1983. The embryonic cell lineage of the nematode Caenorhabditis elegans. Developmental Biology 100: 64–119.CrossRefGoogle ScholarPubMed
Summers, W. C. 2023. The American Phage Group: Founders of Molecular Biology. Yale University Press.Google Scholar
Symonds, N. 1986. What Is Life? Schrödinger’s influence on biology. Quarterly Review of Biology 61: 221–226.CrossRefGoogle ScholarPubMed
Teschendorff, A. E. and Feinberg, A. P. 2021. Statistical mechanics meets single-cell biology. Nature Reviews Genetics 22: 459–476.CrossRefGoogle ScholarPubMed
Thompson, D. W. 1917. On Growth and Form. Cambridge University Press.CrossRefGoogle Scholar
Timoféeff-Ressovsky, N., Zimmer, K., and Delbrück, M. 1935. Über die Natur der Genmutation und der Genstruktur. Nachrichten aus der Biologie der Gesellschaft der Wissenschaften zu Göttingen 1: 189–245.Google Scholar
Tribus, M. and McIrvine, E. C. 1971. Energy and information. Scientific American 225: 179–188.CrossRefGoogle Scholar
Turing, A. 1952. The chemical basis of morphogenesis. Philosophical Transactions of the Royal Society B: Biological Sciences 237: 37–72.Google Scholar
von Foerster, H. 1960. On self-organizing systems and their environments. In Yovits, M. C. and Cameron, S. (eds.), Self-Organizing Systems. Pergamon Press, pp. 31–50.Google Scholar
von Wright, G. H. 1955. Ludwig Wittgenstein, a biographical sketch. Philosophical Review 64: 527–545.CrossRefGoogle Scholar
Waddington, C. H. 1962. New Patterns in Genetics and Development. Columbia University Press.CrossRefGoogle Scholar
Waddington, C. H. 1969. Some European contributions to the prehistory of molecular biology. Nature 221: 318–321.CrossRefGoogle Scholar
Walsby, A. E. and Hodge, M. J. S. 2017. Schrödinger’s code-script: Not a genetic cipher but a code of development. Studies in History and Philosophy of Biological and Biomedical Sciences 63: 45–54.CrossRefGoogle Scholar
Walsh, D. M. 2015. Organism, Agency, and Evolution. Cambridge University Press.CrossRefGoogle Scholar
Walsh, D. M. 2020. Action, program, metaphor. Interdisciplinary Science Reviews 45: 344–359.CrossRefGoogle Scholar
Watson, J. D. 1965. The Molecular Biology of the Gene. W. A. Benjamin, Inc.Google Scholar
Watson, J. D. 1966. Growing up in the phage group. In Cairns, J., Stent, G. S., and Watson, J. D. (eds.), Phage and the Origins of Molecular Biology. Cold Spring Harbor Laboratory Press, pp. 239–245.Google Scholar
Watson, J. D. 1993. Succeeding in science: Some rules of thumb. Science 261: 1812–1813.CrossRefGoogle ScholarPubMed
Watson, J. D. 2001. A Passion for DNA. Cold Spring Harbor Laboratory Press.Google Scholar
Watson, J. D. and Crick, F. H. C. 1953a. Molecular structure of nucleic acids. Nature 171: 737–738.CrossRefGoogle ScholarPubMed
Watson, J. D. and Crick, F. H. C. 1953b. Genetical implications of the structure of deoxyribonucleic acid. Nature 171: 964–967.CrossRefGoogle ScholarPubMed
Weaver, W. 1948. Science and complexity. American Scientist 36: 536–544.Google ScholarPubMed
Weismann, A. 1893. The Germ-Plasm: A Theory of Heredity. Charles Scribner’s Sons.Google Scholar
Weiss, P. A. 1973. The Science of Life: The Living System – A System for Living. Futura.Google Scholar
Wicken, J. 1987. Evolution, Thermodynamics, and Information: Extending the Darwinian Program. Oxford University Press.Google Scholar
Wiener, N. 1948. Cybernetics: Control and Communication in the Animal and the Machine. MIT Press.Google Scholar
Wigner, E. P. and Hodgkin, R. A. 1977. Michael Polanyi (1891–1976). Biographical Memoirs of Fellows of the Royal Society 23: 413–448.Google Scholar
Williams, N. 2016. Irene Manton, Erwin Schrödinger and the puzzle of chromosome structure. Journal of the History of Biology 49: 425–459.CrossRefGoogle ScholarPubMed
Wilkins, M. H. F. 1963. Molecular configuration of nucleic acids. Science 140: 941–950.CrossRefGoogle ScholarPubMed
Windle, B. C. A. 1908. What Is Life? A Study of Vitalism and Neo-Vitalism. Sands.CrossRefGoogle Scholar
Wise, M. N. 1994. Pascual Jordan: Quantum mechanics, psychology, National Socialism. In M. Renneberg and M. Walker (eds.), Science, Technology, and National Socialism. Cambridge University Press, pp. 224–254.Google Scholar
Witkowski, J. A. 1986. Schrödinger’s What Is Life?: Entropy, order, and hereditary code-scripts. Trends in Biochemical Sciences 11: 266–268.CrossRefGoogle Scholar
Woese, C. R. 2004. A new biology for a new century. Microbiology and Molecular Biology Reviews 68: 173–186.CrossRefGoogle ScholarPubMed
Wolpert, L. 1994. Do we understand development? Science 266: 571–572.CrossRefGoogle ScholarPubMed
Wolpert, L. and Lewis, J. H. 1975. Towards a theory of development. In Thorbeke, G. J. (ed.), Biology of Aging and Development. Faseb, pp. 21–34.Google Scholar
Wu, D., Daugherty, S. C., Van Aken, S. E., Pai, G. H., Watkins, K. L., Khouri, H., Tallon, L. J., Zaborsky, J. M., Dunbar, H. E., Tran, P. L., Moran, M. A., and Eisen, K. A. 2006. Metabolic complementarity and genomics of the dual bacterial symbiosis of sharpshooters. PLoS Biology 4: e188.CrossRefGoogle Scholar
Yoxen, E. J. 1977. The social impact of molecular biology. Dissertation, University of Cambridge.Google Scholar
Yoxen, E. J. 1979. Where does Schrödinger’s What Is Life? belong in the history of molecular biology? History of Science 17: 17–52.CrossRefGoogle ScholarPubMed

Accessibility standard: WCAG 2.0 A

Why this information is here

This section outlines the accessibility features of this content - including support for screen readers, full keyboard navigation and high-contrast display options. This may not be relevant for you.

Accessibility Information

The PDF of this Element conforms to version 2.0 of the Web Content Accessibility Guidelines (WCAG), ensuring core accessibility principles are addressed and meets the basic (A) level of WCAG compliance, addressing essential accessibility barriers.

Content Navigation

Table of contents navigation
Allows you to navigate directly to chapters, sections, or non‐text items through a linked table of contents, reducing the need for extensive scrolling.

Reading Order & Textual Equivalents

Single logical reading order
You will encounter all content (including footnotes, captions, etc.) in a clear, sequential flow, making it easier to follow with assistive tools like screen readers.
Short alternative textual descriptions
You get concise descriptions (for images, charts, or media clips), ensuring you do not miss crucial information when visual or audio elements are not accessible.

Structural and Technical Features

ARIA roles provided
You gain clarity from ARIA (Accessible Rich Internet Applications) roles and attributes, as they help assistive technologies interpret how each part of the content functions.

Save element to Kindle

To save this element to your Kindle, first ensure no-reply@cambridge.org is added to your Approved Personal Document E-mail List under your Personal Document Settings on the Manage Your Content and Devices page of your Amazon account. Then enter the ‘name’ part of your Kindle email address below. Find out more about saving to your Kindle.

Note you can select to save to either the @free.kindle.com or @kindle.com variations. ‘@free.kindle.com’ emails are free but can only be saved to your device when it is connected to wi-fi. ‘@kindle.com’ emails can be delivered even when you are not connected to wi-fi, but note that service fees apply.

Find out more about the Kindle Personal Document Service.

What Is Life? Revisited
Available formats
×

Save element to Dropbox

To save content items to your account, please confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your account. Find out more about saving content to Dropbox.

What Is Life? Revisited
Available formats
×

Save element to Google Drive

To save content items to your account, please confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your account. Find out more about saving content to Google Drive.

What Is Life? Revisited
Available formats
×