The Extended Evolutionary Synthesis (EES) consists of a set of theoretical concepts argued to be more comprehensive than the earlier modern synthesis of evolutionary biology that took place between 1918 and 1942. The extended evolutionary synthesis was called for in the 1950s by C. H. Waddington, argued for on the basis of punctuated equilibrium by Stephen Jay Gould and Niles Eldredge in the 1980s, and was reconceptualized in 2007 by Massimo Pigliucci and Gerd B. Müller.

The extended evolutionary synthesis revisits the relative importance of different factors at play, examining several assumptions of the earlier synthesis, and augmenting it with additional causative factors.[1][2] It includes multilevel selection, transgenerational epigenetic inheritance, niche construction, evolvability, and several concepts from evolutionary developmental biology.[3][4][5][6]

Not all biologists have agreed on the need for, or the scope of, an extended synthesis.[7][8][9] Many have collaborated on another synthesis in evolutionary developmental biology, which concentrates on developmental molecular genetics and evolution to understand how natural selection operated on developmental processes and deep homologies between organisms at the level of highly conserved genes.

The preceding "modern synthesis"

Several major ideas about evolution came together in the population genetics of the early 20th century to form the modern synthesis, including genetic variation, natural selection, and particulate (Mendelian) inheritance. This ended the eclipse of Darwinism and supplanted a variety of non-Darwinian theories of evolution. However, it did not unify all of biology, omitting sciences such as developmental biology.

Main article: Modern synthesis (20th century)

The modern synthesis was the widely accepted early-20th-century synthesis reconciling Charles Darwin's theory of evolution by natural selection and Gregor Mendel's theory of genetics in a joint mathematical framework. It established evolution as biology's central paradigm. The 19th-century ideas of natural selection by Darwin and Mendelian genetics were united by researchers who included Ronald Fisher, J. B. S. Haldane and Sewall Wright, the three founders of population genetics, between 1918 and 1932.[10][11][12][13] Julian Huxley introduced the phrase "modern synthesis" in his 1942 book, Evolution: The Modern Synthesis.[14][15][16]

Early history

During the 1950s, English biologist C. H. Waddington called for an extended synthesis based on his research on epigenetics and genetic assimilation.[17][18][19]

In 1978, Michael J. D. White wrote about an extension of the modern synthesis based on new research from speciation.[20] In the 1980s, entomologist Ryuichi Matsuda coined the term "pan-environmentalism" as an extended evolutionary synthesis which he saw as a fusion of Darwinism with neo-Lamarckism.[21] He held that heterochrony is a main mechanism for evolutionary change and that novelty in evolution can be generated by genetic assimilation.[21][22] An extended synthesis was also proposed by the Austrian zoologist Rupert Riedl, with the study of evolvability.[23]

Gordon Rattray Taylor in his 1983 book The Great Evolution Mystery called for an extended synthesis, noting that the modern synthesis is only a subsection of a more comprehensive explanation for biological evolution still to be formulated.[24] In 1985, biologist Robert G. B. Reid authored Evolutionary Theory: The Unfinished Synthesis, which argued that the modern synthesis with its emphasis on natural selection is an incomplete picture of evolution, and emergent evolution can explain the origin of genetic variation.[25][26][27]

In 1988, ethologist John Endler wrote about developing a newer synthesis, discussing processes of evolution that he felt had been neglected.[28]

In 2000, Robert L. Carroll called for an "expanded evolutionary synthesis" due to new research from molecular developmental biology, systematics, geology and the fossil record.[29]

Punctuated equilibrium

In the 1980s, the American palaeontologists Stephen Jay Gould and Niles Eldredge argued for an extended synthesis based on their 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.[30][31][32][33]

Contributions from evolutionary developmental biology

Main article: Evolutionary developmental biology

Some researchers in the field of evolutionary developmental biology proposed another synthesis. They argue that the modern and extended syntheses should mostly center on genes and suggest an integration of embryology with molecular genetics and evolution, aiming to understand how natural selection operates on gene regulation and deep homologies between organisms at the level of highly conserved genes, transcription factors and signalling pathways.[34][5] By contrast, a different strand of evo-devo following an organismal approach[35][36][37][38][39][40] contributes to the extended synthesis by emphasizing (amongst others) developmental bias[41] (both through facilitation[42] and constraint[43]), evolvability,[44][45] and inherency of form [46][47] as primary factors in the evolution of complex structures and phenotypic novelties.[48][49]

Recent history

Massimo Pigliucci, a leading proponent of the extended evolutionary synthesis in its 2007 form

The idea of an extended synthesis was relaunched in 2007 by Massimo Pigliucci,[50][51][52] and Gerd B. Müller,[38][52] with a book in 2010 titled Evolution: The Extended Synthesis, which has served as a launching point for work on the extended synthesis.[52] This includes:

Other processes such as evolvability, phenotypic plasticity, reticulate evolution, horizontal gene transfer, symbiogenesis are said by proponents to have been excluded or missed from the modern synthesis.[59][60] The goal of Piglucci's and Müller's 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.[61]

Michael R. Rose and Todd Oakley have called for a postmodern synthesis, they commented that "it is now abundantly clear that living things often attain a degree of genomic complexity far beyond simple models like the "gene library" genome of the Modern Synthesis".[62] Biologist Eugene Koonin has suggested that the gradualism of the modern synthesis is unsustainable as gene duplication, horizontal gene transfer and endosymbiosis play a pivotal role in evolution.[63] Koonin commented 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."[63]

Arlin Stoltzfus and colleagues advocate mutational and developmental bias in the introduction of variation as an important source of orientation or direction in evolutionary change.[64][65][66][67] They argue that bias in the introduction of variation was not formally recognized throughout the 20th century, due to the influence of neo-Darwinism on thinking about causation.[68]

Organism-centered evolution

The early biologists of the organicist movement have influenced the modern extended evolutionary synthesis. Recent research has called for expanding the population genetic framework of evolutionary biology by a more organism-centered perspective.[69][70] This has been described as "organism-centered evolution" which looks beyond the genome to the ways that individual organisms are participants in their own evolution.[70][71][72] Philip Ball has written a research review on organism-centered evolution.[73][74]

Rui Diogo has proposed a revision of evolutionary theory, which he has termed ONCE: Organic Nonoptimal Constrained Evolution.[75] According to ONCE, evolution is mainly driven by the behavioural choices and persistence of organisms themselves, whilst natural selection plays a secondary role.[75][76][77] ONCE cites examples of reciprocal causation between organism and the environment, Baldwin effect, organic selection, developmental bias and niche construction.[76][77][78]


The extended synthesis is characterized by its additional set of predictions that differ from the standard modern synthesis theory:

  1. Change in phenotype can precede change in genotype[4]
  2. Changes in phenotype are predominantly positive, rather than neutral (see: neutral theory of molecular evolution)
  3. Changes in phenotype are induced in many organisms, rather than one organism[4]
  4. Revolutionary change in phenotype can occur through mutation, facilitated variation[4] or threshold events[49][79]
  5. Repeated evolution in isolated populations can be by convergent evolution or developmental bias[4][41]
  6. Adaptation can be caused by natural selection, environmental induction, non-genetic inheritance, learning and cultural transmission (see: Baldwin effect, meme, transgenerational epigenetic inheritance, ecological inheritance, non-Mendelian inheritance)[4]
  7. Rapid evolution can result from simultaneous induction, natural selection[4] and developmental dynamics[80]
  8. Biodiversity can be affected by features of developmental systems such as differences in evolvability[4]
  9. Heritable variation is directed towards variants that are adaptive and integrated with phenotype[4]
  10. Niche construction is biased towards environmental changes that suit the constructor's phenotype, or that of its descendants, and enhance their fitness[2]
  11. Kin selection[3]
  12. Multilevel selection[4]
  13. Self-organization[50][81]
  14. Symbiogenesis[60][82][83]


From 2016 to 2019, there was an organized project entitled "Putting The Extended Evolutionary Synthesis To The Test" supported by a 7.5 million USD grant from the John Templeton Foundation, supplemented with further money from participating instutitions including Clark University, Indiana University, Lund University, Stanford University, University of Southampton and University of St Andrews.[84]

Publications from the project include over 200 papers, a special issue,[85] and an anthology on Evolutionary Causation.[86] In 2019 a final report of the 2016–2019 consortium was published, Putting the Extended Evolutionary Synthesis to the Test.[87]

The project was headed by Kevin N. Laland at the University of St Andrews and Tobias Uller at Lund University. According to Laland what the extended synthesis "really boils down to is recognition that, in addition to selection, drift, mutation and other established evolutionary processes, other factors, particularly developmental influences, shape the evolutionary process in important ways."[88]


Biologists disagree on the need for an extended synthesis. Opponents contend that the modern synthesis is able to fully account for the newer observations, whereas others criticize the extended synthesis for not being radical enough.[89] Proponents think that the conceptions of evolution at the core of the modern synthesis are too narrow[90] and 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 distorts the picture of biology that modern experimentation has discovered.[91] 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][92] In 2022, the John Templeton Foundation published a review of recent literature.[93]


  1. ^ Wade, Michael J. (2011). "The Neo-Modern Synthesis: The Confluence of New Data and Explanatory Concepts". BioScience. 61 (5): 407–408. doi:10.1525/bio.2011.61.5.10.
  2. ^ a b c d Laland, Kevin N.; Uller, Tobias; Feldman, Marcus W.; Sterelny, Kim; Müller, Gerd B.; Moczek, Armin; Jablonka, Eva; Odling-Smee, John (2015-08-22). "The extended evolutionary synthesis: its structure, assumptions and predictions". Proc. R. Soc. B. 282 (1813): 20151019. doi:10.1098/rspb.2015.1019. PMC 4632619. PMID 26246559.
  3. ^ a b Danchin, É.; Charmantier, A.; Champagne, F. A.; Mesoudi, A.; Pujol, B.; Blanchet, S (2011). "Beyond DNA: integrating inclusive inheritance into an extended theory of evolution". Nature Reviews Genetics. 12 (7): 475–486. doi:10.1038/nrg3028. PMID 21681209. S2CID 8837202.
  4. ^ a b c d e f g h i j Pigliucci, Massimo; Finkelman, Leonard (2014). "The Extended (Evolutionary) Synthesis Debate: Where Science Meets Philosophy". BioScience. 64 (6): 511–516. doi:10.1093/biosci/biu062.
  5. ^ a b c 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 (7): 565–577. doi:10.1002/jez.b.22631. PMC 4744698. PMID 26097188.
  6. ^ Müller, Gerd B. (December 2007). "Evo–devo: extending the evolutionary synthesis". Nature Reviews Genetics. 8 (12): 943–949. doi:10.1038/nrg2219. PMID 17984972. S2CID 19264907.
  7. ^ Svensson, E.I. (2023). The Structure of Evolutionary Theory: Beyond Neo-Darwinism, Neo-Lamarckism and Biased Historical Narratives About the Modern Synthesis. In: Dickins, T.E., Dickins, B.J. (eds) Evolutionary Biology: Contemporary and Historical Reflections Upon Core Theory. Springer. pp. 173–217. ISBN 978-3-031-22027-2
  8. ^ Charlesworth D, Barton NH, Charlesworth B. (2017). "The sources of adaptive variation". Proceedings of the Royal Society B. 284 (1855): 20162864. doi:10.1098/rspb.2016.2864. PMC 5454256. PMID 28566483.((cite journal)): CS1 maint: multiple names: authors list (link)
  9. ^ Futuyma, Douglas J. (2017). "Evolutionary biology today and the call for an extended synthesis". Interface Focus. 7: 20160145. doi:10.1098/rsfs.2016.0145. PMC 5566807.
  10. ^ National Academy of Sciences (1999). Science and Creationism: A View from the National Academy of Sciences (2nd ed.). Washington, D.C.: National Academy Press. p. 28. doi:10.17226/6024. ISBN 978-0-309-06406-4. LCCN 99006259. OCLC 43803228. PMID 25101403. The scientific consensus around evolution is overwhelming.
  11. ^ Bock, Walter J. (July 1981). "Reviewed Work: The Evolutionary Synthesis. Perspectives on the Unification of Biology". The Auk. 98 (3): 644–646. ISSN 0004-8038. JSTOR 4086148.
  12. ^ Fisher, Ronald A. (January 1919). "XV.—The Correlation between Relatives on the Supposition of Mendelian Inheritance". Transactions of the Royal Society of Edinburgh. 52 (2): 399–433. doi:10.1017/S0080456800012163. ISSN 0080-4568. OCLC 4981124. S2CID 181213898. "Paper read by J. Arthur Thomson on July 8, 1918 to the Royal Society of Edinburgh."
  13. ^ Fisher, R. A. (1999) [Originally published 1930; Oxford, UK: The Clarendon Press]. The Genetical Theory of Natural Selection. Edited with a foreword and notes by J. H. Bennett (A complete variorum ed.). Oxford, UK: Oxford University Press. ISBN 978-0-19-850440-5. LCCN 00702764. OCLC 45308589.
  14. ^ Hubbs, C.L. (1943). "Evolution the new synthesis". American Naturalist. 77 (771): 365–68. doi:10.1086/281134.
  15. ^ Kimball, R.F. (1943). "The great biological generalization". Quarterly Review of Biology. 18 (4): 364–67. doi:10.1086/394682. S2CID 88212178.
  16. ^ Karl P. Schmidt, Evolution the Modern Synthesis by Julian Huxley, Copeia, Vol. 1943, No. 4 (Dec. 31, 1943), pp. 262-263
  17. ^ Wilkins, Adam S (2008). "Waddington's Unfinished Critique of Neo-Darwinian Genetics: Then and Now". Biological Theory. 3 (3): 224–232. doi:10.1162/biot.2008.3.3.224. S2CID 84217300.
  18. ^ Pigliucci, Massimo; et al. (2006). "Phenotypic plasticity and evolution by genetic assimilation". Journal of Experimental Biology. 209 (12): 2362–2367. doi:10.1242/jeb.02070. PMID 16731812.
  19. ^ Huang, Sui (2012). "The molecular and mathematical basis of Waddington's epigenetic landscape: A framework for post-Darwinian biology?". BioEssays. 34 (2): 149–157. doi:10.1002/bies.201100031. ISSN 0265-9247. PMID 22102361. S2CID 19632484.
  20. ^ Parnell, Dennis R. (1978). "Heralding a New Synthesis: Modes of Speciation by M. J. D. White". Systematic Botany. 3 (1): 126. doi:10.2307/2418537. JSTOR 2418537.
  21. ^ a b Pearson, Roy Douglas (1988). "Animal Evolution in Changing Environments". Acta Biotheoretica. 37: 31–36. doi:10.1007/BF00050806.
  22. ^ Shapiro, Arthur M. (1988). "Animal Evolution in Changing Environments" (PDF). Journal of the Lepidopterists' Society. 42 (2): 146–147. Archived from the original on 2023-11-05. Retrieved 2023-11-05.((cite journal)): CS1 maint: bot: original URL status unknown (link)
  23. ^ Wagner, Günter P; Laubichler; Manfred D. (2004). "Rupert Riedl and the Re-Synthesis of Evolutionary and Developmental Biology: Body Plans and Evolvability" Archived 2015-12-08 at the Wayback Machine. Journal of Experimental Zoology (Mol Dev Evol) 302B: 92-102.
  24. ^ Birx, H. James (1984). "Neo-Darwinism and Neo-Social Darwinism". BioScience. 34 (3): 196–197. doi:10.2307/1309778.
  25. ^ Hoff, Charles (1986). "Evolutionary Theory: The Unfinished Synthesis by Robert G. B. Reid". Human Biology. 58 (5): 823–824. JSTOR 41463815.
  26. ^ William, Mary B. (1986). "Evolutionary Theory: The Unfinished Synthesis by Robert G. B. Reid". The Quarterly Review of Biology. 61 (2): 266. doi:10.1086/414957. JSTOR 2829141.
  27. ^ Cornell, John F. (1987). "Evolutionary Theory: The Unfinished Synthesis by Robert G. B. Reid". Journal of the History of Biology. 20 (3): 424–425. JSTOR 4331027.
  28. ^ Endler, John A; McLellan, Tracy (1988). "The Processes of Evolution: Toward a Newer Synthesis". Annual Review of Ecology and Systematics. 19: 395–421. doi:10.1146/ JSTOR 2097160.
  29. ^ Carroll, Robert L. (2000). "Towards a new evolutionary synthesis". Trends in Ecology & Evolution. 15 (1): 27–32. doi:10.1016/s0169-5347(99)01743-7. PMID 10603504.
  30. ^ Gould, Stephen Jay. (1980). Is a New and General Theory of Evolution Emerging? Paleobiology. Vol. 6, No. 1. pp. 119-130.
  31. ^ Gould, Stephen Jay (1982). "Darwinism and the Expansion of Evolutionary Theory". Science. 216 (4544): 380–387. Bibcode:1982Sci...216..380G. doi:10.1126/science.7041256. PMID 7041256.
  32. ^ "A More Modern Synthesis". American Scientist.
  33. ^ Vermeij, Geerat J (1987). "Unfinished Synthesis: Biological Hierarchies and Modern Evolutionary Thought by Niles Eldredge". The Quarterly Review of Biology. 62 (1): 79–80. doi:10.1086/415312.
  34. ^ Davidson, Eric H. (2006). The regulatory genome : gene regulatory networks in development and evolution. Amsterdam [Netherlands]. ISBN 978-0120885633. OCLC 61756485.((cite book)): CS1 maint: location missing publisher (link)
  35. ^ Bateson, P (2005). "The Return of the Whole Organism". Journal of Biosciences. 30 (1): 31–39. doi:10.1007/bf02705148. PMID 15824439. S2CID 26656790.
  36. ^ Huneman, Philippe (2010). "Assessing the Prospects for a Return of Organisms in Evolutionary Biology". History and Philosophy of the Life Sciences. 32 (2–3): 341–372. PMID 21162374.
  37. ^ Gilbert, S.F.; Opitz, G.; Raff, R. (1996). "Resynthesizing Evolutionary and Developmental Biology". Developmental Biology. 173 (2): 357–372. doi:10.1006/dbio.1996.0032. PMID 8605997.
  38. ^ a b Müller, G. B. (2007). "Evo-devo: Extending the evolutionary synthesis". Nature Reviews Genetics. 8 (12): 943–949. doi:10.1038/nrg2219. PMID 17984972. S2CID 19264907.
  39. ^ "The Origins of Form". Natural History.
  40. ^ Hall, Brian K. (1998). Evolutionary developmental biology (2nd ed.). London: Chapman & Hall. ISBN 978-0412785801. OCLC 40606316.
  41. ^ a b c Brakefield, Paul M. (July 2006). "Evo-devo and constraints on selection". Trends in Ecology & Evolution. 21 (7): 362–368. doi:10.1016/j.tree.2006.05.001. ISSN 0169-5347. PMID 16713653.
  42. ^ Gerhart, John; Kirschner, Marc (2007-05-15). "The theory of facilitated variation". Proceedings of the National Academy of Sciences. 104 (suppl 1): 8582–8589. Bibcode:2007PNAS..104.8582G. doi:10.1073/pnas.0701035104. PMC 1876433. PMID 17494755.
  43. ^ a b Smith, J. Maynard; Burian, R.; Kauffman, S.; Alberch, P.; Campbell, J.; Goodwin, B.; Lande, R.; Raup, D.; Wolpert, L. (September 1985). "Developmental Constraints and Evolution: A Perspective from the Mountain Lake Conference on Development and Evolution". The Quarterly Review of Biology. 60 (3): 265–287. doi:10.1086/414425. ISSN 0033-5770. S2CID 85201850.
  44. ^ Wagner, Günter P.; Altenberg, Lee (June 1996). "Perspective: Complex Adaptations and the Evolution of Evolvability". Evolution. 50 (3): 967–976. doi:10.1111/j.1558-5646.1996.tb02339.x. ISSN 0014-3820. PMID 28565291. S2CID 21040413.
  45. ^ Huneman, Philippe; Walsh, Denis M. (2017-08-17). Challenging the modern synthesis : adaptation, development, and inheritance. Huneman, Philippe,, Walsh, Denis M., 1958-. New York, NY. ISBN 9780199377183. OCLC 1001337947.((cite book)): CS1 maint: location missing publisher (link)
  46. ^ Newman, Stuart A.; Müller, Gerd B. (2006-01-06), "Genes and Form", Genes in Development, Duke University Press, pp. 38–73, doi:10.1215/9780822387336-003, ISBN 9780822387336
  47. ^ Newman, Stuart A. (2017-11-15), "Inherency", Evolutionary Developmental Biology, Springer International Publishing, pp. 1–12, doi:10.1007/978-3-319-33038-9_78-1, ISBN 9783319330389
  48. ^ a b Müller, Gerd B. (2010-03-26), "Epigenetic Innovation", Evolution—the Extended Synthesis, The MIT Press, pp. 307–333, doi:10.7551/mitpress/9780262513678.003.0012, ISBN 9780262513678, retrieved 2018-06-22
  49. ^ a b c Peterson, Tim; Müller, Gerd B. (2016-04-28). "Phenotypic Novelty in EvoDevo: The Distinction Between Continuous and Discontinuous Variation and Its Importance in Evolutionary Theory". Evolutionary Biology. 43 (3): 314–335. doi:10.1007/s11692-016-9372-9. ISSN 0071-3260. PMC 4960286. PMID 27512237.
  50. ^ a b 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. S2CID 2703146.
  51. ^ Grant, Bob (1 January 2010). "Should Evolutionary Theory Evolve". The Scientist.
  52. ^ a b c d e Pigliucci, Massimo; Müller, Gerd B., eds. (26 March 2010). Evolution - the Extended Synthesis. The MIT Press. ISBN 978-0262513678.
  53. ^ Meaden, Rhiannon (5 August 2015). "Redefining Evolutionary Biolog". The Royal Society Publishing Blog. Archived from the original on 23 October 2015. Retrieved 29 September 2015.
  54. ^ Indiana University (7 August 2015). "Expanding the Theory of Evolution". Lab Manager.
  55. ^ Bonduriansky, R; Day, T (2009). "Nongenetic inheritance and its evolutionary implications" (PDF). Annual Review of Ecology and Systematics. 40: 103–125. doi:10.1146/annurev.ecolsys.39.110707.173441.
  56. ^ 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 3335599. PMID 22567381.
  57. ^ 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 4138557. PMID 25191292.
  58. ^ Moczek, Armin P. (2011-05-05). "Evolutionary biology: The origins of novelty". Nature. 473 (7345): 34–35. Bibcode:2011Natur.473...34M. doi:10.1038/473034a. PMID 21544136. S2CID 4413435.
  59. ^ Perez, JUlio E; Alfonsi, Carmen; Munoz, Carlos (2010). "Towards a New Evolutionary Theory" (PDF). Interciencia. 35: 862–868.
  60. ^ a b 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
  61. ^ "Expanding Theory of Evolution". PhysOrg. 5 August 2015.
  62. ^ Rose, Michael R.; Oakley, Todd H. (November 24, 2007). "The new biology: beyond the Modern Synthesis" (PDF). Biology Direct. 2 (30): 30. doi:10.1186/1745-6150-2-30. PMC 2222615. PMID 18036242. Archived (PDF) from the original on 2014-03-21.
  63. ^ a b Koonin, Eugene (2009). "Towards a postmodern synthesis of evolutionary biology". Cell Cycle. 8 (6): 799–800. doi:10.4161/cc.8.6.8187. PMC 3410441. PMID 19242109.
  64. ^ Yampolsky, L. Y.; Stoltzfus, A. (2001). "Bias in the introduction of variation as an orienting factor in evolution". Evolution & Development. 3 (2): 73–83. doi:10.1046/j.1525-142x.2001.003002073.x. PMID 11341676. S2CID 26956345.
  65. ^ Stoltzfus, A. (2006). "Mutation-Biased Adaptation in a Protein NK Model". Molecular Biology and Evolution. 23 (10): 1852–1862. doi:10.1093/molbev/msl064. PMID 16857856.
  66. ^ Stoltzfus, A.; Yampolsky, L. Y. (2009). "Climbing Mount Probable: Mutation as a Cause of Nonrandomness in Evolution". Journal of Heredity. 100 (5): 637–647. doi:10.1093/jhered/esp048. PMID 19625453.
  67. ^ Stoltzfus, Arlin (2017). "Why we don't want another "Synthesis"". Biology Direct. 12 (1): 23. doi:10.1186/s13062-017-0194-1. PMC 5625744. PMID 28969666.
  68. ^ A. Stoltzfus (2021). Mutation, Randomness and Evolution. Oxford, Oxford.
  69. ^ Nicholson, Daniel J. (2014). "The Return of the Organism as a Fundamental Explanatory Concept in Biology". Philosophy Compass. 9 (5): 347–359. doi:10.1111/phc3.12128.
  70. ^ a b Baedke, J., Fábregas-Tejeda, A. (2023). The Organism in Evolutionary Explanation: From Early Twentieth Century to the Extended Evolutionary Synthesis. In: Dickins, T.E., Dickins, B.J. (eds) Evolutionary Biology: Contemporary and Historical Reflections Upon Core Theory. Springer. pp. 121–150. ISBN 978-3031220272
  71. ^ "What Is Organism-Centered Evolution?". Retrieved 4 November 2023.
  72. ^ "Extended Evolutionary Synthesis: A review of the latest scientific research". Retrieved 4 November 2023.
  73. ^ "Organisms as Agents of Evolution: New Research Review". Retrieved 4 November 2023.
  74. ^ "Organisms as Agents of Evolution". Retrieved 4 November 2023.
  75. ^ a b Smulders, Tom V. (2017). "Evolution Driven by Organismal Behaviour – a Unifying View of Life, Function, Form, Mismatches, and Trends". Journal of Anatomy. 232 (2): 356–357. doi:10.1111/joa.12750. PMC 5770302.
  76. ^ a b Fleagle, John G. (2017). "Evolution Driven by Organismal Behavior: A Unifying View of Life, Function, Form, Mismatches, and Trends". The Quarterly Review of Biology. 92 (4): 469. doi:10.1086/694961.
  77. ^ a b "Evolution Driven by Organismal Behavior". Retrieved 10 November 2023.
  78. ^ Sánchez-Villagra, Marcelo R. (2018). "Evolution Driven by Organismal Behavior: A Unifying View of Life, Function, Form, Mismatches and Trends". Swiss Journal of Palaeontology. 137: 109–112. doi:10.1007/s13358-017-0139-4.
  79. ^ Lange, Axel; Nemeschkal, Hans L.; Müller, Gerd B. (April 2018). "A threshold model for polydactyly". Progress in Biophysics and Molecular Biology. 137: 1–11. doi:10.1016/j.pbiomolbio.2018.04.007. ISSN 0079-6107. PMID 29739620.
  80. ^ Favé, Marie-Julie; Johnson, Robert A.; Cover, Stefan; Handschuh, Stephan; Metscher, Brian D.; Müller, Gerd B.; Gopalan, Shyamalika; Abouheif, Ehab (2015-09-04). "Past climate change on Sky Islands drives novelty in a core developmental gene network and its phenotype". BMC Evolutionary Biology. 15 (1): 183. doi:10.1186/s12862-015-0448-4. ISSN 1471-2148. PMC 4560157. PMID 26338531.
  81. ^ Johnson, BR; Lam, SK (2010). "Self-Organization, Natural Selection, and Evolution: Cellular Hardware and Genetic Software". BioScience. 60 (11): 879–885. doi:10.1525/bio.2010.60.11.4. S2CID 10903076.
  82. ^ Gontier, Nathalie. (2016). History of Symbiogenesis. In Richard M Kliman. Encyclopedia of Evolutionary Biology. Elsevier Science. pp. 261-271. ISBN 978-0128000496
  83. ^ Agafonov VA, Negrobov VV, Igamberdiev AU. (2021). "Symbiogenesis as a driving force of evolution: The legacy of Boris Kozo Polyansky". Biosystems. 199: 104302. doi:10.1016/j.biosystems.2020.104302. PMID 33227379.((cite journal)): CS1 maint: multiple names: authors list (link)
  84. ^ "Should Evolutionary Theory Evolve". University of Southampton. 7 April 2016.
  85. ^ "Our special issue on Developmental Bias in Evolution is officially published online – Extended Evolutionary Synthesis". Retrieved 2023-01-27.
  86. ^ "Evolutionary Causation". MIT Press. Retrieved 2023-01-27.
  87. ^ "Putting the Extended Evolutionary Synthesis to the Test. Retrieved 22 August 2022.
  88. ^ "Empowering the Extended Evolutionary Synthesis". The Evolution Institute.
  89. ^ Craig, Lindsay R. (June 2010). "The So-Called Extended Synthesis and Population Genetics". Biological Theory. 5 (2): 117–123. doi:10.1162/biot_a_00035. ISSN 1555-5542. S2CID 84662773.
  90. ^ 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.((cite journal)): CS1 maint: multiple names: authors list (link)
  91. ^ 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.
  92. ^ Müller, Gerd B. (2017-10-06). "Why an extended evolutionary synthesis is necessary". Interface Focus. 7 (5): 20170015. doi:10.1098/rsfs.2017.0015. ISSN 2042-8898. PMC 5566817. PMID 28839929.
  93. ^ "Extended Evolutionary Synthesis". John Templeton Foundation. Retrieved 2023-01-27.

Further reading

Defence of the extended synthesis

Criticism of the extended synthesis