Biological Theory

, Volume 7, Issue 4, pp 298–310 | Cite as

Formalization and the Meaning of “Theory” in the Inexact Biological Sciences

  • James Griesemer
Thematic Issue Article: The Meaning of "Theory" in Biology


Exact sciences are described as sciences whose theories are formalized. These are contrasted to inexact sciences, whose theories are not formalized. Formalization is described as a broader category than mathematization, involving any form/content distinction allowing forms, e.g., as represented in theoretical models, to be studied independently of the empirical content of a subject-matter domain. Exactness is a practice depending on the use of theories to control subject-matter domains and to align theoretical with empirical models and not merely a state of a science. Inexact biological sciences tolerate a degree of “mismatch” between theoretical and empirical models and concepts. Three illustrations from biological sciences are discussed in which formalization is achieved by various means: Mendelism, Weismannism, and Darwinism. Frege’s idea of a “conceptual notation” is used to further characterize the notion of a form/content distinction.


Darwin Exact and inexact science Formalization Mendel, model Theory Weismann 



Thanks are due to workshop organizers Massimo Pigliucci, Werner Callebaut, and Kim Sterelny, workshop participants, and the Konrad Lorenz Institute for Evolution and Cognition Research. I thank Roberta Millstein, Elihu Gerson, and the workshop organizers for valuable comments on a previous version of the manuscript and Ken Waters for encouragement. Support of a UC Davis Herbert A. Young Society Deans’ Fellowship, 2011–2014, is gratefully acknowledged.


  1. Brandon R (1990) Adaptation and environment. Princeton University Press, PrincetonGoogle Scholar
  2. Brigandt I (2010) Beyond reduction and pluralism: toward an epistemology of explanatory integration in biology. Erkenntnis 73:295–311CrossRefGoogle Scholar
  3. Bulmer M (1985) The mathematical theory of quantitative genetics. Oxford University Press, New YorkGoogle Scholar
  4. Buss L (1987) The evolution of individuality. Princeton University Press, PrincetonGoogle Scholar
  5. Calcott B, Sterelny K (eds) (2011) The major transitions in evolution revisited. MIT Press, CambridgeGoogle Scholar
  6. Darwin C ([1838] 1987) E Notebook. In: Barrett P, Gautrey P, Herbert S, Kohn D, Smith S (eds) Charles Darwin’s notebooks, 1836–1844: geology, transmutation of species, metaphysical enquiries. Cambridge University Press, Cambridge, pp 395–455Google Scholar
  7. Darwin C ([1859] 1964) On the origin of species. Facsimile of the 1st edition. Harvard University Press, Cambridge, MAGoogle Scholar
  8. Dawkins R (1982) The extended phenotype. Oxford University Press, New YorkGoogle Scholar
  9. Eldredge N, Cracraft J (1980) Phylogenetic patterns and the evolutionary process: method and theory in comparative biology. Columbia University Press, New YorkGoogle Scholar
  10. Felsenstein J (2004) Inferring phylogenies. Sinauer, SunderlandGoogle Scholar
  11. Frege G ([1879] 1972) Begriffschrift. Translated as “Conceptual notation”. In: Bynum TW (trans) Conceptual notation and related articles. Clarendon Press, Oxford, pp 101–203Google Scholar
  12. Frege G ([1882a] 1972) Über die wissenschaftliche Berechtigung einer Begriffsschrift. Translated as “On the scientific justification of a conceptual notation”. In: Bynum TW (trans) Conceptual notation and related articles. Clarendon Press, Oxford, pp 83–89Google Scholar
  13. Frege G ([1882b] 1972) “Über den Zweck der Begriffsschrift,” translated as “On the aim of the ‘Conceptual Notation.’” In: Bynum TW (trans) Conceptual notation and related articles. Clarendon Press, Oxford, pp 90–100Google Scholar
  14. Galton F (1876) A theory of heredity. J Anthropol Inst Great Brit Ireland 5:329–348CrossRefGoogle Scholar
  15. Gánti T (2003) The principles of life, with a commentary by James Griesemer and Eörs Szathmáry. Vekerdi L, Czaran E, Muller V, trans. Oxford University Press, OxfordGoogle Scholar
  16. Giere RN (1988) Explaining science: a cognitive approach. University of Chicago Press, ChicagoCrossRefGoogle Scholar
  17. Giere RN (2006) Scientific perspectivism. University of Chicago Press, ChicagoCrossRefGoogle Scholar
  18. Giere RN, Bickle J, Mauldin R (2006) Understanding scientific reasoning, 5th edn. Wadsworth, BelmontGoogle Scholar
  19. Gilbert SF (1985) Developmental biology, 1st edn. Sinauer, SunderlandGoogle Scholar
  20. Gillespie J (2004) Population genetics: a concise guide, 2nd edn. Johns Hopkins University Press, BaltimoreGoogle Scholar
  21. Godfrey-Smith P (2009) Darwinian populations and natural selection. Oxford University Press, New YorkGoogle Scholar
  22. Goodman N (1976) Languages of art, 2nd edn. Hackett, IndianapolisGoogle Scholar
  23. Griesemer J (1984) Presentations and the status of theories. In: Asquith PD, Kitcher P (eds) PSA 1984, vol. 1. Philosophy of Science Association, East Lansing, MI, pp 102–114Google Scholar
  24. Griesemer J (1990) Modeling in the museum: on the role of remnant models in the work of Joseph Grinnell. Biol Philos 5:3–36CrossRefGoogle Scholar
  25. Griesemer J (2000) Development, culture and the units of inheritance. Philos Sci 67:S348–S368CrossRefGoogle Scholar
  26. Griesemer J (2007) Tracking organic processes: representations and research styles in classical embryology and genetics. In: Laubichler MD, Maienschein J (eds) From embryology to EvoDevo: a history of developmental evolution. MIT Press, Cambridge, pp 375–433Google Scholar
  27. Griesemer J (2011a) Heuristic reductionism and the relative significance of epigenetic inheritance in evolution. In: Hallgrímsson B, Hall BK (eds) Epigenetics: linking genotype and phenotype in development and evolution. University of California Press, Los Angeles, pp 14–40Google Scholar
  28. Griesemer J (2011b) The relative significance of epigenetic inheritance in evolution: some philosophical considerations. In: Gissis S, Jablonka E (eds) Transformations of Lamarckism: from subtle fluids to molecular biology. MIT Press, Cambridge, pp 331–344Google Scholar
  29. Griesemer J, Wimsatt WC (1989) Picturing Weismannism: a case study of conceptual Evolution. In: Ruse M (ed) What the philosophy of biology is: essays for David Hull. Kluwer, Dordrecht, pp 75–137CrossRefGoogle Scholar
  30. Hagen J (2003) The statistical frame of mind in systematic biology from quantitative zoology to biometry. J Hist Biol 36:353–384CrossRefGoogle Scholar
  31. Hastings A (1997) Population biology: concepts and models. Springer, New YorkCrossRefGoogle Scholar
  32. Hull DL (1988) Science as a process. University of Chicago Press, ChicagoGoogle Scholar
  33. Kuhn TS (1970) The structure of scientific revolutions, 2nd edn. University of Chicago Press, ChicagoGoogle Scholar
  34. Levins R (1966) The strategy of model building in population biology. Am Sci 54:421–431Google Scholar
  35. Lewontin RC (1970) The units of selection. Annu Rev Ecol Syst 1:1–17CrossRefGoogle Scholar
  36. Lloyd EA (1994) The structure and confirmation of evolutionary theory. Princeton University Press, PrincetonGoogle Scholar
  37. Love AC (2008) Explaining evolutionary innovation and novelty: criteria of adequacy and multidisciplinary prerequisites. Philos Sci 75:874–886CrossRefGoogle Scholar
  38. Love AC (2012) Theory is as theory does: scientific practice and theory structure in biology. Biol Theory 7. doi: 10.1007/s13752-012-0046-2
  39. Maienschein J (1978) Cell lineage, ancestral reminiscence, and the biogenetic law. J Hist Biol 11:129–158CrossRefGoogle Scholar
  40. Maynard Smith J (1965) The theory of evolution, 2nd edn. Penguin, LondonGoogle Scholar
  41. Maynard Smith J, Szathmáry E (1995) The major transitions in evolution. Freeman (Spektrum), OxfordGoogle Scholar
  42. Mendel G ([1866] 1965) Experiments in plant hybridization. Royal Horticultural Society of London (trans). Harvard University Press, Cambridge, MAGoogle Scholar
  43. Morgan M, Morrison M (1999) Models as mediators: perspectives on natural and social science. Cambridge University Press, CambridgeCrossRefGoogle Scholar
  44. Morrison M (2007) Where have all the theories gone? Philos Sci 74:195–228CrossRefGoogle Scholar
  45. Morrison M (2011) One phenomenon, many models: inconsistency and complementarity. Stud Hist Philos Sci 42:342–351CrossRefGoogle Scholar
  46. Nagel E, Newman JR (1958) Gödel’s proof. New York University Press, New YorkGoogle Scholar
  47. Nogler G (2006) The lesser-known Mendel: his experiments on Hieracium. Genetics 172:1–6Google Scholar
  48. Perini L (2004) The truth in pictures. Philos Sci 72:262–285CrossRefGoogle Scholar
  49. Slatkin M (2009) Epigenetic inheritance and the missing heritability problem. Genetics 182:845–850CrossRefGoogle Scholar
  50. Suppes P (1960) A comparison of the meaning and uses of models in mathematics and the empirical sciences. Synthese 12:287–301CrossRefGoogle Scholar
  51. Suppes P (1962) Models of data. In: Nagel E, Suppes P, Tarski A (eds) Logic, methodology and philosophy of science. Stanford University Press, Stanford, pp 252–261Google Scholar
  52. Suppes P (1967) What is a scientific theory? In: Morgenbesser S (ed) Philosophy of science today. Basic Books, New York, pp 55–67Google Scholar
  53. Tal O, Kisdi E, Jablonka E (2010) Epigenetic contribution to covariance between relatives. Genetics 184:1037–1050CrossRefGoogle Scholar
  54. Teller P (2001) Twilight of the perfect model model. Erkenntnis 55:393–415CrossRefGoogle Scholar
  55. Thompson DW (2000) On growth and form. Cambridge University Press (Canto), New YorkGoogle Scholar
  56. Thompson P (2007) Formalisations of evolutionary biology. In: Matthen M, Stephens C (eds) Philosophy of biology. Elsevier, Amsterdam, pp 485–523CrossRefGoogle Scholar
  57. van Fraassen B (1980) The scientific image. Clarendon Press, OxfordCrossRefGoogle Scholar
  58. Wagner G (2010) The measurement theory of fitness. Evolution 64:1358–1376Google Scholar
  59. Walter H (1913) Genetics. MacMillan, New YorkGoogle Scholar
  60. Waters CK (2008) How practical know-how contextualizes theoretical knowledge: exporting causal knowledge from laboratory to nature. Philos Sci 75:707–719CrossRefGoogle Scholar
  61. Weismann A (1893) The Germ-plasm, a theory of heredity. In: Parker W, Ronnfeldt H (trans). Scribner’s, New York. Orig. Das Keimplasma, eine theorie der Vererbung, Jena, Gustav FischerGoogle Scholar
  62. Whitman CO (1887) A contribution to the history of the germ-layers in clepsine. J Morphol 1:105–182CrossRefGoogle Scholar
  63. Wilson EB (1896) The cell in development and inheritance. MacMillan, LondonCrossRefGoogle Scholar
  64. Wimsatt WC (1980) Reductionistic research strategies and their biases in the units of selection controversy. In: Nickles T (ed) Scientific discovery, vol II., Historical and scientific case studies Reidel, Dordrecth, pp 213–259CrossRefGoogle Scholar
  65. Wimsatt WC (1987) False models as means to truer theories. In: Nitecki M, Hoffman A (eds) Neutral models in biology. Oxford University Press, London, pp 23–55Google Scholar
  66. Wimsatt WC (2007) Re-engineering philosophy for limited beings: piecewise approximations to reality. Harvard University Press, CambridgeGoogle Scholar
  67. Wimsatt WC, Griesemer J (2007) Reproducing entrenchments to scaffold culture: the central role of development in cultural evolution. In: Sansom R, Brandon R (eds) Integrating evolution and development: from theory to practice. MIT Press, Cambridge, pp 227–323Google Scholar
  68. Winsberg E (2006) Handshaking your way to the top: simulation at the nanoscale. Philos Sci 73:582–594CrossRefGoogle Scholar
  69. Winsberg E (2010) Science in the age of computer simulation. University of Chicago Press, ChicagoCrossRefGoogle Scholar

Copyright information

© Konrad Lorenz Institute for Evolution and Cognition Research 2012

Authors and Affiliations

  1. 1.Department of PhilosophyUniversity of California, DavisDavisUSA

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