Extended Evolutionary Synthesis

Massimo Pigliucci, a leading proponent of the extended evolutionary synthesis.

The extended evolutionary synthesis is an extension of the mid-20th century Modern Synthesis of evolutionary biology. It revisits the relative importance of different factors at play, revisits several assumptions of the original synthesis, and augments it with additional causative factors in evolution.[1][2]

The extended synthesis includes concepts and mechanisms such as multilevel selection theory, transgenerational epigenetic inheritance, niche construction and evolvability.[3][4][5]

Early history

During the 1950s, biologist C. H. Waddington called for an extended synthesis based from his research on epigenetics and genetic assimilation.[6][7][8] An extended synthesis was also proposed by Rupert Riedl, with the study of evolvability.[9] In 1978, Michael J. D. White wrote about an extension of the modern synthesis based on new research from speciation.[10]

In the 1980s, Stephen Jay Gould argued for an extended synthesis. This was based on his idea of punctuated equilibrium, the role of species selection shaping large scale evolutionary patterns and natural selection working on multiple levels extending from genes to species.[11][12][13] Similar ideas were held by Niles Eldredge author of the book Unfinished Synthesis (1985).[14]

Ethologist John Endler wrote a paper in 1988 discussing processes of evolution that he felt had been neglected.[15] Others, such as researchers in the field of evolutionary developmental biology (evo-devo) called for a different extended synthesis based on the opinion that the modern synthesis was mostly centered on genes and ignored the development of the organism, as well as morphological areas of biology.[16][17][18][19][20]

Recent history

The extended synthesis is still a work-in-progress, essentially having been launched in 2007 by a paper by Massimo Pigliucci in the journal Evolution,[21] followed by a conference on the subject in 2008 at the Konrad Lorenz Institute for Evolution and Cognition Research.[22] The participants of the conference published a book in 2010 titled Evolution: The Extended Synthesis, which has served as a launching point for work on the extended synthesis.[23] Several of the topics included in the extended synthesis include:

Other processes such as evolvability, phenotypic plasticity, reticulate evolution, sex evolution[30] and symbiogenesis are said by proponents to have been excluded or missed from the modern synthesis.[31][32]

The goal of the extended synthesis is to take evolution beyond the gene-centered approach of population genetics to consider more organism- and ecology-centered approaches. Many of these causes are currently considered secondary in evolutionary causation, and proponents of the extended synthesis want them to be considered first-class evolutionary causes.[33]

Biologist Eugene Koonin writing on the extended synthesis in 2009 wrote that "the new developments in evolutionary biology by no account should be viewed as refutation of Darwin. On the contrary, they are widening the trails that Darwin blazed 150 years ago and reveal the extraordinary fertility of his thinking."[34]


The Extended Evolutionary Synthesis proposed by Pigliucci and colleagues makes the following predictions:

  1. change in phenotype can precede change in genotype
  2. changes in phenotype are predominantly positive, rather than neutral (see: neutral theory of molecular evolution)
  3. changes in phentotype are induced in many organisms, rather than one organism
  4. revolutionary change in phenotype can occur through mutation or facilitated variation
  5. "repeated evolution in isolated populations may be due to convergent selection and/or developmental bias"
  6. adaptation occur due to natural selection, environmental induction, non-genetic inheritance, learning and cultural transmission (see: Baldwin effect, meme, transgenerational epigenetic inheritance, ecological inheritance, non-Mendelian inheritance)
  7. rapid evolution can result from the simultaneous induction and natural selection
  8. biodiversity can be affected by features of developmental systems (evolvability, constraints)
  9. heritable variation is directed towards variants that are adaptive and integrated with phenotype
  10. "niche construction will be systematically biased towards environmental changes that are well suited to the constructor's phenotype, or that of its descendants, and enhance the constructor's, or its descendant's, fitness"[35]
  11. multilevel selection[4]
  12. kin selection[3]


Biologists disagree on the need for an extended synthesis. Opponents contend that the modern synthesis is able to fully account for the newer observations, while proponents think that the conceptions of evolution at the core of the modern synthesis are too narrow.[36] Proponents argue that even when the modern synthesis allows for the ideas in the extended synthesis, using the modern synthesis affects the way that biologists think about evolution. For example, Denis Noble says that using terms and categories of the modern synthesis distort the picture of biology that modern experimentation has discovered.[37] Proponents therefore claim that the extended synthesis is necessary to help expand the conceptions and framework of how evolution is considered throughout the biological disciplines.[2]

The ideas of the extended synthesis were positively reviewed by Anya Plutynski for the National Center for Science Education.[38]

Proponents of Extended Evolutionary Synthesis state that there are many adaptation mechanisms, while most of biologist maintain that there is only one - natural selection.[39]


  1. Wade, Michael J. (2011). "The Neo-Modern Synthesis: The Confluence of New Data and Explanatory Concepts". BioScience 61: 407-408.
  2. 1 2 John Odling-Smee et al. "The extended evolutionary synthesis: its structure, assumptions and predictions". Proceedings of the Royal Society B: Biological Sciences, August 2015.
  3. 1 2 Danchin É, Charmantier A, Champagne FA, Mesoudi A, Pujol B, Blanchet S. (2011). "Beyond DNA: integrating inclusive inheritance into an extended theory of evolution". Nature Reviews Genetics 12: 475-486.
  4. 1 2 Pigliucci, Massimo; Finkelman, Leonard. "The Extended (Evolutionary) Synthesis Debate: Where Science Meets Philosophy". BioScience 64: 511-516, 2014.
  5. 1 2 Laubichler, Manfred D; Renn, Jürgen. (2015). "Extended evolution: A Conceptual Framework for Integrating Regulatory Networks and Niche Construction". Journal of Experimental Zoology Part B: Molecular and Developmental Evolution 324: 565–577.
  6. Wilkins, Adam S. (2008). Waddington's Unfinished Critique of Neo-Darwinian Genetics: Then and Now. Biological Theory 3 (3):224-232.
  7. Massimo Pigliucci et al. (2006). "Phenotypic plasticity and evolution by genetic assimilation". Journal of Experimental Biology 209: 2362-2367.
  8. Huang S. (2011). The molecular and mathematical basis of Waddington’s epigenetic landscape: A framework for post-Darwinian biology? BioEssays 34: 149-157.
  9. Wagner, Günter P; Laubichler; Manfred D. (2004). "Rupert Riedl and the Re-Synthesis of Evolutionary and Developmental Biology: Body Plans and Evolvability". Journal of Experimental Zoology (Mol Dev Evol) 302B: 92-102.
  10. Parnell, Dennis R. (1978). Heralding a New Synthesis Modes of Speciation by M. J. D. White. Systematic Botany. Vol. 3, No. 1. p. 126.
  11. Gould, Stephen Jay. (1980). Is a New and General Theory of Evolution Emerging? Paleobiology. Vol. 6, No. 1. pp. 119-130.
  12. Gould, Stephen Jay. (1982). Darwinism and the Expansion of Evolutionary Theory. Science. New Series, Vol. 216, No. 4544. pp. 380-387.
  13. "A More Modern Synthesis". American Scientist.
  14. Vermeij, Geerat J. (1987). Unfinished Synthesis: Biological Hierarchies and Modern Evolutionary Thought by Niles Eldredge. The Quarterly Review of Biology. Vol. 62, No. 1. pp. 79-80.
  15. Endler, John A; McLellan, Tracy. (1988). The Processes of Evolution: Toward a Newer Synthesis. Annual Review of Ecology and Systematics. Vol. 19, pp. 395-421.
  16. Bateson P. (2005). The Return of the Whole Organism. Journal of Biosciences 30: 31-39.
  17. Huneman, Philippe. (2010). Assessing the Prospects for a Return of Organisms in Evolutionary Biology. History and Philosophy of the Life Sciences 32: 341-372.
  18. Gilbert S.F., Opitz G., Raff R. (1996). Resynthesizing Evolutionary and Developmental Biology. Development and Evolution 173: 357-372.
  19. Müller, G. B. (2007). Evo-devo: Extending the evolutionary synthesis. Nature Reviews Genetics 8: 943-949.
  20. "The Origins of Form". Natural History.
  21. Pigliucci, Massimo (2007). "Do We Need an Extended Evolutionary Synthesis?". Evolution. 61 (12): 2743–2749. doi:10.1111/j.1558-5646.2007.00246.x. PMID 17924956.
  22. Grant, Bob (1 January 2010). "Should Evolutionary Theory Evolve". The Scientist.
  23. 1 2 3 Pigliucci (26 March 2010). Evolution - the Extended Synthesis. The MIT Press. ISBN 978-0262513678.
  24. Meaden, Rhiannon (5 August 2015). "Redefining Evolutionary Biology". The Royal Society Publishing Blog.
  25. Indiana University (7 August 2015). "Expanding the Theory of Evolution". Lab Manager.
  26. Bonduriansky, R; Day, T. (2009). "Nongenetic inheritance and its evolutionary implications". Annual Review of Ecology and Systematics 40: 103-125.
  27. Schrey; et al. (15 December 2011). "The Role of Epigenetic in Evolution: the Extended Synthesis". Genetics Research International. 2012: 286164. doi:10.1155/2012/286164. PMC 3335599Freely accessible. PMID 22567381.
  28. Stotz, Karola (20 August 2014). "Extended evolutionary psychology: the importance of transgenerational developmental plasticity". Frontiers in Psychology. 5: 908. doi:10.3389/fpsyg.2014.00908. PMC 4138557Freely accessible. PMID 25191292.
  29. Laland; et al. (5 August 2015). "The extended evolutionary synthesis: its structure, assumptions and predictions". The Royal Society Publishing Proceedings B. 282 (1813): 20151019. doi:10.1098/rspb.2015.1019. PMID 26246559.
  30. Th Lodé 2011. Sex is not a solution for reproduction, the libertine bubble theory. Bioessays 33:419-422
  31. Perez, JUlio E; Alfonsi, Carmen; Munoz, Carlos. (2010). "Towards a New Evolutionary Theory". Interciencia 35: 862-868.
  32. Gontier, Nathalie. (2015). Reticulate Evolution Everywhere. In Reticulate Evolution: Symbiogenesis, Lateral Gene Transfer, Hybridization and Infectious Heredity. Springer. pp. 1-40. ISBN 978-3-319-16344-4
  33. "Expanding Theory of Evolution". PhysOrg. 5 August 2015.
  34. Koonin, Eugene. (2009). "Towards a postmodern synthesis of evolutionary biology". Cell Cycle 8: 799-800.
  35. The extended evolutionary synthesis: its structure, assumptions and predictions, Proc Biol Sci. 2015 Aug 22; 282(1813): 20151019. doi: 10.1098/rspb.2015.1019
  36. Laland, Kevin, Tobias Uller, Marc Feldman, Kim Sterelny, Gerd B. Müller, Armin Moczek, Eva Jablonka, John Odling-Smee, Gregory A. Wray, Hopi E. Hoekstra, Douglas J. Futuyma, Richard E. Lenski, Trudy F. C. Mackay, Dolph Schluter & Joan E. Strassmann; et al. (8 October 2014). "Does Evolutionary Theory Need a Rethink?". Nature. 514 (7521): 161–164. Bibcode:2014Natur.514..161L. doi:10.1038/514161a. PMID 25297418.
  37. Noble, Denis (1 January 2015). "Evolution Beyond Neo-Darwinism: A New Conceptual Framework". The Journal of Experimental Biology. 218 (Pt 1): 7–13. doi:10.1242/jeb.106310. PMID 25568446.
  38. "Evolution: The Extended Synthesis". National Center for Science Education.
  39. Scott-Phillips, T. C., Laland, K. N., Shuker, D. M., Dickins, T. E. and West, S. A. (2014). "The Niche Construction Perspective: A Critical Appraisal". Evolution 68: 1231-1243.

Further reading

Defend the extended synthesis

Criticism of the extended synthesis

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