Extended evolutionary synthesis

File:Modern Synthesis.svg 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|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.{{cite book |author=National Academy of Sciences |author-link=National Academy of Sciences |year=1999 |title=Science and Creationism: A View from the National Academy of Sciences |url=https://archive.org/details/sciencecreationi0000unse/page/28 |edition=2nd |location=Washington, D.C. |publisher=National Academy Press |isbn=978-0-309-06406-4 |lccn=99006259 |oclc=43803228 |page=[https://archive.org/details/sciencecreationi0000unse/page/28 28] |quote=The scientific consensus around evolution is overwhelming. |doi=10.17226/6024 |pmid=25101403 }}{{cite journal |last=Bock |first=Walter J. |date=July 1981 |title=Reviewed Work: The Evolutionary Synthesis. Perspectives on the Unification of Biology |journal=The Auk |volume=98 |issue=3 |pages=644–646 |issn=0004-8038 |jstor=4086148}}{{cite journal |last=Fisher |first=Ronald A. |author-link=Ronald Fisher |date=January 1919 |title=XV.—The Correlation between Relatives on the Supposition of Mendelian Inheritance |journal=Transactions of the Royal Society of Edinburgh |volume=52 |issue=2 |pages=399–433 |doi=10.1017/S0080456800012163 |s2cid=181213898 |issn=0080-4568 |oclc=4981124|url=https://zenodo.org/record/1428666 }} "Paper read by J. Arthur Thomson on July 8, 1918 to the Royal Society of Edinburgh."{{cite book |last=Fisher |first=R. A. |author-link=Ronald Fisher |year=1999 |orig-year=Originally published 1930; Oxford, UK: The Clarendon Press |title=The Genetical Theory of Natural Selection |others=Edited with a foreword and notes by J. H. Bennett |edition=A complete variorum |location=Oxford, UK |publisher=Oxford University Press |isbn=978-0-19-850440-5 |lccn=00702764 |oclc=45308589|title-link=The Genetical Theory of Natural Selection }} Julian Huxley introduced the phrase "modern synthesis" in his 1942 book, Evolution: The Modern Synthesis.{{cite journal | last1 = Hubbs | first1 = C.L. | year = 1943 | title = Evolution the new synthesis | journal = American Naturalist | volume = 77 | issue = 771| pages = 365–68 | doi=10.1086/281134}}{{cite journal | last1 = Kimball | first1 = R.F. | year = 1943 | title = The great biological generalization | journal = Quarterly Review of Biology | volume = 18 | issue = 4| pages = 364–67 | doi=10.1086/394682| s2cid = 88212178 }}Karl P. Schmidt, Evolution the Modern Synthesis by Julian Huxley, Copeia, Vol. 1943, No. 4 (Dec. 31, 1943), pp. 262-263

During the 1950s, English biologist C. H. Waddington called for an extended synthesis based on his research on epigenetics and genetic assimilation.{{cite journal | last1=Wilkins | first1=Adam S | s2cid=84217300 | year=2008 | title=Waddington's Unfinished Critique of Neo-Darwinian Genetics: Then and Now | journal=Biological Theory | volume=3 | issue=3| pages=224–232 | doi=10.1162/biot.2008.3.3.224}}{{cite journal | last1=Pigliucci | first1=Massimo | author-link=Massimo Pigliucci | display-authors=etal | year=2006 | title=Phenotypic plasticity and evolution by genetic assimilation | journal=Journal of Experimental Biology | volume=209 | issue=12| pages=2362–2367 | doi=10.1242/jeb.02070| pmid=16731812 | doi-access=free}}{{cite journal |last=Huang |first=Sui |title=The molecular and mathematical basis of Waddington's epigenetic landscape: A framework for post-Darwinian biology? |journal=BioEssays |volume=34 |issue=2 |year=2012 |pages=149–157 |issn=0265-9247 |doi=10.1002/bies.201100031|pmid=22102361 |s2cid=19632484 }}

In 1978, Michael J. D. White wrote about an extension of the modern synthesis based on new research from speciation.{{cite journal|author=Parnell, Dennis R.|year=1978|title=Heralding a New Synthesis: Modes of Speciation by M. J. D. White|journal=Systematic Botany|volume=3|issue=1|pages=126|doi=10.2307/2418537|jstor=2418537}} 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.{{cite journal|author=Pearson, Roy Douglas|year=1988|title=Animal Evolution in Changing Environments|journal=Acta Biotheoretica|volume=37|issue=|pages=31–36|doi=10.1007/BF00050806}} He held that heterochrony is a main mechanism for evolutionary change and that novelty in evolution can be generated by genetic assimilation.{{cite journal|author=Shapiro, Arthur M.|year=1988|title=Animal Evolution in Changing Environments|journal=Journal of the Lepidopterists' Society|url=https://images.peabody.yale.edu/lepsoc/jls/1980s/1988/1988-42%282%29146-Shapiro.pdf|volume=42|issue=2|pages=146–147|access-date=2023-11-05|archive-date=2023-11-05|archive-url=https://web.archive.org/web/20231105175658/https://images.peabody.yale.edu/lepsoc/jls/1980s/1988/1988-42%282%29146-Shapiro.pdf|url-status=bot: unknown}} An extended synthesis was also proposed by the Austrian zoologist Rupert Riedl, with the study of evolvability.Wagner, Günter P; Laubichler; Manfred D. (2004). [http://www.yale.edu/gpwagner/pdfs/Riedl2004.pdf "Rupert Riedl and the Re-Synthesis of Evolutionary and Developmental Biology: Body Plans and Evolvability"] {{webarchive|url=https://web.archive.org/web/20151208094623/http://www.yale.edu/gpwagner/pdfs/Riedl2004.pdf |date=2015-12-08 }}. Journal of Experimental Zoology (Mol Dev Evol) 302B: 92-102.

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.{{cite journal|author=Birx, H. James|year=1984|title=Neo-Darwinism and Neo-Social Darwinism|journal= BioScience|volume=34|issue=3|pages=196–197|doi=10.2307/1309778|jstor=1309778 }} 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.{{cite journal|author=Hoff, Charles|year=1986|title=Evolutionary Theory: The Unfinished Synthesis by Robert G. B. Reid|journal=Human Biology|volume=58|issue=5|pages=823–824 |jstor=41463815}}{{cite journal|author=William, Mary B.|year=1986|title=Evolutionary Theory: The Unfinished Synthesis by Robert G. B. Reid|journal=The Quarterly Review of Biology|url=https://www.journals.uchicago.edu/doi/10.1086/414957|volume=61|issue=2|pages=266|doi=10.1086/414957| jstor=2829141}}{{cite journal|author=Cornell, John F.|year=1987|title=Evolutionary Theory: The Unfinished Synthesis by Robert G. B. Reid|journal=Journal of the History of Biology|volume=20|issue=3|pages=424–425|jstor=4331027}}

In 1988, ethologist John Endler wrote about developing a newer synthesis, discussing processes of evolution that he felt had been neglected.{{cite journal | last1 = Endler | first1 = John A | last2 = McLellan | first2 = Tracy | year = 1988 | title = The Processes of Evolution: Toward a Newer Synthesis | jstor = 2097160 | journal = Annual Review of Ecology and Systematics | volume = 19 | pages = 395–421 | doi = 10.1146/annurev.es.19.110188.002143 }}

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.{{cite journal|author=Carroll, Robert L.|year=2000|title=Towards a new evolutionary synthesis|journal=Trends in Ecology & Evolution|url=https://www.sciencedirect.com/science/article/abs/pii/S0169534799017437|volume=15|issue=1|pages=27–32|doi=10.1016/s0169-5347(99)01743-7|pmid=10603504}}

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.Gould, Stephen Jay. (1980). Is a New and General Theory of Evolution Emerging? Paleobiology. Vol. 6, No. 1. pp. 119-130.{{cite journal | last1=Gould | first1=Stephen Jay | year=1982 | title=Darwinism and the Expansion of Evolutionary Theory | journal=Science | volume=216 | issue=4544| pages=380–387 | doi=10.1126/science.7041256| pmid=7041256 | bibcode=1982Sci...216..380G}}[http://www.americanscientist.org/bookshelf/pub/a-more-modern-synthesis "A More Modern Synthesis"]. American Scientist.{{cite journal | last1=Vermeij | first1=Geerat J | year=1987 | title=Unfinished Synthesis: Biological Hierarchies and Modern Evolutionary Thought by Niles Eldredge | journal=The Quarterly Review of Biology | volume=62 | issue=1| pages=79–80 | doi=10.1086/415312}}

{{main|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.{{Cite book|title=The regulatory genome : gene regulatory networks in development and evolution|author=Davidson, Eric H.|isbn=978-0120885633|location=Amsterdam [Netherlands]|oclc=61756485|year = 2006}} By contrast, a different strand of evo-devo following an organismal approach{{cite journal | last1=Bateson | first1=P | s2cid=26656790 | year=2005 | title=The Return of the Whole Organism | journal=Journal of Biosciences | volume=30 | issue=1| pages=31–39 | doi=10.1007/bf02705148 | pmid=15824439}}{{cite journal | last1=Huneman | first1=Philippe | year=2010 | title=Assessing the Prospects for a Return of Organisms in Evolutionary Biology | journal=History and Philosophy of the Life Sciences | volume=32 | issue=2–3 | pages=341–372 | pmid=21162374 }}{{cite journal | last1=Gilbert | first1=S.F. | last2=Opitz | first2=G. | last3=Raff | first3=R. | year=1996 | title=Resynthesizing Evolutionary and Developmental Biology | journal= Developmental Biology| volume=173 | issue=2| pages=357–372 | doi=10.1006/dbio.1996.0032 | pmid=8605997| doi-access=free }}{{cite journal | last1=Müller | first1=G. B. | s2cid=19264907 | author-link=Gerd B. Müller | year=2007 | title=Evo-devo: Extending the evolutionary synthesis | journal=Nature Reviews Genetics | volume=8 | issue=12 | pages=943–949 | doi=10.1038/nrg2219 | pmid=17984972}}[http://www.naturalhistorymag.com/features/061488/the-origins-of-form "The Origins of Form"]. Natural History.{{Cite book|title=Evolutionary developmental biology|author=Hall, Brian K. |date=1998|publisher=Chapman & Hall|isbn=978-0412785801|edition= 2nd|location=London|oclc=40606316}} contributes to the extended synthesis by emphasizing (amongst others) developmental bias{{Cite journal|last=Brakefield|first=Paul M.|date=July 2006|title=Evo-devo and constraints on selection|journal=Trends in Ecology & Evolution|volume=21|issue=7|pages=362–368|doi=10.1016/j.tree.2006.05.001|pmid=16713653|issn=0169-5347}} (both through facilitation{{Cite journal|last1=Gerhart|first1=John|last2=Kirschner|first2=Marc|date=2007-05-15|title=The theory of facilitated variation|journal=Proceedings of the National Academy of Sciences|volume=104|issue=suppl 1|pages=8582–8589|doi=10.1073/pnas.0701035104|pmc=1876433|pmid=17494755|bibcode=2007PNAS..104.8582G|doi-access=free}} and constraint{{Cite journal|last1=Smith|first1=J. Maynard|last2=Burian|first2=R.|last3=Kauffman|first3=S.|last4=Alberch|first4=P.|last5=Campbell|first5=J.|last6=Goodwin|first6=B.|last7=Lande|first7=R.|last8=Raup|first8=D.|last9=Wolpert|first9=L.|date=September 1985|title=Developmental Constraints and Evolution: A Perspective from the Mountain Lake Conference on Development and Evolution|journal=The Quarterly Review of Biology|volume=60|issue=3|pages=265–287|doi=10.1086/414425|s2cid=85201850|issn=0033-5770}}), evolvability,{{Cite journal|last1=Wagner|first1=Günter P.|last2=Altenberg|first2=Lee|s2cid=21040413|date=June 1996|journal=Evolution|volume=50|issue=3|pages=967–976|doi=10.1111/j.1558-5646.1996.tb02339.x|pmid=28565291|issn=0014-3820|title=Perspective: Complex Adaptations and the Evolution of Evolvability|doi-access=free}}{{Cite book|title=Challenging the modern synthesis : adaptation, development, and inheritance|others=Huneman, Philippe,, Walsh, Denis M., 1958-|isbn=9780199377183|location=New York, NY|oclc=1001337947|last1 = Huneman|first1 = Philippe|last2=Walsh|first2=Denis M.|date=2017-08-17}} and inherency of form {{Citation|last1=Newman|first1=Stuart A.|title=Genes and Form|date=2006-01-06|work=Genes in Development|pages=38–73|publisher=Duke University Press|doi=10.1215/9780822387336-003|isbn=9780822387336|last2=Müller|first2=Gerd B.}}{{Citation|last=Newman|first=Stuart A.|chapter=Inherency|date=2017-11-15|pages=1–12|publisher=Springer International Publishing|doi=10.1007/978-3-319-33038-9_78-1|isbn=9783319330389|title=Evolutionary Developmental Biology|doi-access=free}} as primary factors in the evolution of complex structures and phenotypic novelties.{{Citation|last=Müller|first=Gerd B.|chapter=Epigenetic Innovation|date=2010-03-26|chapter-url=http://mitpress.universitypressscholarship.com/view/10.7551/mitpress/9780262513678.001.0001/upso-9780262513678-chapter-12|pages=307–333|publisher=The MIT Press|doi=10.7551/mitpress/9780262513678.003.0012|isbn=9780262513678|access-date=2018-06-22|title=Evolution—the Extended Synthesis}}{{Cite journal|last1=Peterson|first1=Tim|last2=Müller|first2=Gerd B.|date=2016-04-28|title=Phenotypic Novelty in EvoDevo: The Distinction Between Continuous and Discontinuous Variation and Its Importance in Evolutionary Theory|journal=Evolutionary Biology|volume=43|issue=3|pages=314–335|doi=10.1007/s11692-016-9372-9|issn=0071-3260|pmc=4960286|pmid=27512237}}

File:20110409-542-NECSS2011.jpg, 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,{{cite journal |last1=Pigliucci |first1=Massimo |s2cid=2703146 | author1-link=Massimo Pigliucci |title=Do We Need an Extended Evolutionary Synthesis? |journal=Evolution |volume=61|issue=12 |pages=2743–2749 |doi=10.1111/j.1558-5646.2007.00246.x |pmid=17924956 |year=2007|url=https://philpapers.org/rec/PIGDWN |doi-access=free }}{{cite web |last1=Grant|first1=Bob|title=Should Evolutionary Theory Evolve|url=http://www.the-scientist.com/?articles.view/articleNo/27894/title/Should-Evolutionary-Theory-Evolve-/|website=The Scientist|date=1 January 2010}} and Gerd B. Müller, with a book in 2010 titled Evolution: The Extended Synthesis, which has served as a launching point for work on the extended synthesis.{{cite book |editor1=Pigliucci, Massimo | editor1-link=Massimo Pigliucci |editor2=Müller, Gerd B. |title=Evolution - the Extended Synthesis |date=26 March 2010 |publisher=The MIT Press |isbn=978-0262513678 |url=http://muse.jhu.edu/books/9780262315142}} This includes:

• The role of prior configurations, genomic structures, and other traits in the organism in generating evolutionary variations.{{cite web |last1=Meaden |first1=Rhiannon |title=Redefining Evolutionary Biolog |work=The Royal Society Publishing Blog |date=5 August 2015 |url=https://blogs.royalsociety.org/publishing/redefining-evolutionary-biology/ |access-date=29 September 2015 |archive-date=23 October 2015 |archive-url=https://web.archive.org/web/20151023182538/https://blogs.royalsociety.org/publishing/redefining-evolutionary-biology/ |url-status=dead }}{{cite web |last1=Indiana University |title=Expanding the Theory of Evolution |url=http://www.labmanager.com/news/2015/08/expanding-the-theory-of-evolution |website=Lab Manager |date=7 August 2015}}
• How increasing dimensionality of fitness landscapes affects our view of speciation.
• The role of multilevel selection in the major evolutionary transitions.
• New types of inheritance, including cultural and epigenetic inheritance.{{cite journal | last1=Bonduriansky | first1=R | last2=Day | first2=T | year=2009 | title=Nongenetic inheritance and its evolutionary implications | url=http://bonduriansky.net/AREES-2009.pdf | journal=Annual Review of Ecology and Systematics | volume=40 | pages=103–125 | doi=10.1146/annurev.ecolsys.39.110707.173441}}{{cite journal |last1=Schrey |title=The Role of Epigenetic in Evolution: the Extended Synthesis |journal=Genetics Research International |date=15 December 2011 |volume=2012 |pages=286164 |doi=10.1155/2012/286164 |pmid=22567381 |pmc=3335599 |display-authors=etal|doi-access=free }}
• The way that organismal development and developmental plasticity channel evolutionary pathways{{cite journal|last1=Stotz |first1=Karola |title=Extended evolutionary psychology: the importance of transgenerational developmental plasticity|journal=Frontiers in Psychology |date=20 August 2014|volume=5|page=908 |doi=10.3389/fpsyg.2014.00908 |pmid=25191292 |pmc=4138557|doi-access=free }} and generates phenotypic novelty{{Cite journal |last=Moczek |first=Armin P. |s2cid=4413435 |date=2011-05-05|title=Evolutionary biology: The origins of novelty |journal=Nature |volume=473|issue=7345 |pages=34–35|doi=10.1038/473034a |pmid=21544136|bibcode=2011Natur.473...34M |doi-access=free }}
• How organisms modify the environments they belong to through niche construction.

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.{{cite journal | last1=Perez | first1=JUlio E | last2=Alfonsi | first2=Carmen | last3=Munoz | first3=Carlos | year=2010 | title=Towards a New Evolutionary Theory | url=http://www.redalyc.org/pdf/339/33915598013.pdf | journal=Interciencia | volume=35 | pages=862–868 }}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}} 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.{{cite news |title=Expanding Theory of Evolution |url=http://phys.org/news/2015-08-theory-evolution.html |publisher=PhysOrg |date=5 August 2015}}

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".{{cite journal |last1=Rose |first1=Michael R. |author-link1=Michael R. Rose |last2=Oakley |first2=Todd H. |date=November 24, 2007 |title=The new biology: beyond the Modern Synthesis |url=http://www.biologydirect.com/content/pdf/1745-6150-2-30.pdf |archive-url=https://web.archive.org/web/20140321054654/http://www.biologydirect.com/content/pdf/1745-6150-2-30.pdf |archive-date=2014-03-21 |url-status=live |journal=Biology Direct |volume=2 |issue=30 |pages=30 |doi=10.1186/1745-6150-2-30 |pmc=2222615 |pmid=18036242 |doi-access=free}} 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.{{cite journal | last1=Koonin | first1=Eugene | year=2009 | title=Towards a postmodern synthesis of evolutionary biology | journal=Cell Cycle | volume=8 | issue=6| pages=799–800 | pmc=3410441 | pmid=19242109 | doi=10.4161/cc.8.6.8187}} 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."

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.{{cite journal |last1=Yampolsky |first1=L. Y. |last2=Stoltzfus |first2=A. |year=2001 |title=Bias in the introduction of variation as an orienting factor in evolution |journal=Evolution & Development |volume=3 | pages=73–83 |doi=10.1046/j.1525-142x.2001.003002073.x |pmid=11341676 |issue=2|s2cid=26956345 }}{{cite journal | last=Stoltzfus |first=A. | title=Mutation-Biased Adaptation in a Protein NK Model | journal=Molecular Biology and Evolution | volume=23 | issue=10 | pages=1852–1862 | year=2006 | doi=10.1093/molbev/msl064 | pmid=16857856 | doi-access=free }}{{cite journal |last1=Stoltzfus |first1=A. |last2=Yampolsky |first2=L. Y. |year=2009 |title=Climbing Mount Probable: Mutation as a Cause of Nonrandomness in Evolution |journal=Journal of Heredity |volume=100 | pages=637–647 |doi=10.1093/jhered/esp048 |pmid=19625453 |issue=5 |doi-access=free}}{{cite journal|author=Stoltzfus, Arlin|year=2017|title=Why we don't want another "Synthesis"|journal=Biology Direct|volume=12|issue=1|pages=23|doi=10.1186/s13062-017-0194-1|pmid=28969666|doi-access=free|pmc=5625744}} 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.{{cite book | author=A. Stoltzfus | year=2021 | title=Mutation, Randomness and Evolution | publisher=Oxford, Oxford }}

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.{{cite journal |author=Nicholson, Daniel J.| year=2014 | title=The Return of the Organism as a Fundamental Explanatory Concept in Biology | journal=Philosophy Compass |url=https://compass.onlinelibrary.wiley.com/doi/abs/10.1111/phc3.12128 |volume=9| issue=5| pages= 347–359| doi=10.1111/phc3.12128}}{{cite book | chapter-url=https://link.springer.com/chapter/10.1007/978-3-031-22028-9_8 | doi=10.1007/978-3-031-22028-9_8 | chapter=The Organism in Evolutionary Explanation: From Early Twentieth Century to the Extended Evolutionary Synthesis | title=Evolutionary Biology: Contemporary and Historical Reflections Upon Core Theory | series=Evolutionary Biology – New Perspectives on Its Development | date=2023 | last1=Baedke | first1=Jan | last2=Fábregas-Tejeda | first2=Alejandro | volume=6 | pages=121–150 | isbn=978-3-031-22027-2 }} 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.[https://web.archive.org/web/20230523164933/https://www.templeton.org/news/what-is-organism-centered-evolution "What Is Organism-Centered Evolution?"]. templeton.org. Retrieved 4 November 2023.[https://web.archive.org/web/20231104232740/https://www.templeton.org/wp-content/uploads/2022/08/EES_Review_FINAL_.pdf "Extended Evolutionary Synthesis: A review of the latest scientific research"]. templeton.org. Retrieved 4 November 2023. Philip Ball has written a research review on organism-centered evolution.[https://www.templeton.org/news/organisms-as-agents-of-evolution-new-research-review "Organisms as Agents of Evolution: New Research Review"]. templeton.org. Retrieved 4 November 2023.[https://www.templeton.org/wp-content/uploads/2023/04/Biological-Agency_1_FINAL.pdf "Organisms as Agents of Evolution"]. templeton.org. Retrieved 4 November 2023.

Rui Diogo has proposed a revision of evolutionary theory, which he has termed ONCE: Organic Nonoptimal Constrained Evolution.{{cite journal|author=Smulders, Tom V.|year=2017|title= Evolution Driven by Organismal Behaviour – a Unifying View of Life, Function, Form, Mismatches, and Trends|journal=Journal of Anatomy|volume=232|issue=2|pages=356–357|doi=10.1111/joa.12750|doi-access=free|pmc=5770302}} According to ONCE, evolution is mainly driven by the behavioural choices and persistence of organisms themselves, whilst natural selection plays a secondary role.{{cite journal|author=Fleagle, John G.|year=2017|title=Evolution Driven by Organismal Behavior: A Unifying View of Life, Function, Form, Mismatches, and Trends |journal=The Quarterly Review of Biology|url=https://www.journals.uchicago.edu/doi/10.1086/694961|volume=92|issue=4|pages=469|doi=10.1086/694961}}[https://extendedevolutionarysynthesis.com/evolution-driven-by-organismal-behavior/ "Evolution Driven by Organismal Behavior"]. extendedevolutionarysynthesis.com. Retrieved 10 November 2023. ONCE cites examples of reciprocal causation between organism and the environment, Baldwin effect, organic selection, developmental bias and niche construction.{{cite journal|author=Sánchez-Villagra, Marcelo R.|year=2018|title=Evolution Driven by Organismal Behavior: A Unifying View of Life, Function, Form, Mismatches and Trends |journal=Swiss Journal of Palaeontology|url=https://sjpp.springeropen.com/articles/10.1007/s13358-017-0139-4|volume=137|issue=|pages=109–112|doi=10.1007/s13358-017-0139-4}}

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
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. Revolutionary change in phenotype can occur through mutation, facilitated variation or threshold events{{Cite journal|last1=Lange|first1=Axel|last2=Nemeschkal|first2=Hans L.|last3=Müller|first3=Gerd B.|date=April 2018|title=A threshold model for polydactyly|journal=Progress in Biophysics and Molecular Biology|volume=137|pages=1–11|doi=10.1016/j.pbiomolbio.2018.04.007|pmid=29739620|issn=0079-6107|doi-access=free}}
5. Repeated evolution in isolated populations can be by convergent evolution or developmental bias
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)
7. Rapid evolution can result from simultaneous induction, natural selection and developmental dynamics{{Cite journal|last1=Favé|first1=Marie-Julie|last2=Johnson |first2=Robert A. |last3=Cover |first3=Stefan |last4=Handschuh |first4=Stephan |last5=Metscher |first5=Brian D. |last6=Müller |first6=Gerd B. |last7=Gopalan |first7=Shyamalika |last8=Abouheif |first8=Ehab |date=2015-09-04 |title=Past climate change on Sky Islands drives novelty in a core developmental gene network and its phenotype|journal=BMC Evolutionary Biology |volume=15|issue=1|pages=183 |doi=10.1186/s12862-015-0448-4|issn=1471-2148|pmc=4560157|pmid=26338531 |doi-access=free }}
8. Biodiversity can be affected by features of developmental systems such as differences in evolvability
9. Heritable variation is directed towards variants that are adaptive and integrated with phenotype
10. Niche construction is biased towards environmental changes that suit the constructor's phenotype, or that of its descendants, and enhance their fitness
11. Kin selection
12. Multilevel selection
13. Self-organization{{cite journal | last1 = Johnson | first1 = BR | last2 = Lam | first2 = SK | s2cid = 10903076 | year = 2010 | title = Self-Organization, Natural Selection, and Evolution: Cellular Hardware and Genetic Software | url = https://academic.oup.com/bioscience/article/60/11/879/328810 | journal = BioScience | volume = 60 | issue = 11| pages = 879–885 | doi=10.1525/bio.2010.60.11.4}}
14. SymbiogenesisGontier, Nathalie. (2016). [https://www.sciencedirect.com/science/article/abs/pii/B9780128000496000160 History of Symbiogenesis]. In Richard M Kliman. Encyclopedia of Evolutionary Biology. Elsevier Science. pp. 261-271. {{ISBN|978-0128000496}}{{cite journal|author=Agafonov VA, Negrobov VV, Igamberdiev AU.|year=2021|title=Symbiogenesis as a driving force of evolution: The legacy of Boris Kozo Polyansky|journal=Biosystems|volume=199|issue=|pages=104302|doi=10.1016/j.biosystems.2020.104302|pmid=33227379}}

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.{{cite web |title=Should Evolutionary Theory Evolve |url=https://www.southampton.ac.uk/news/2016/04/evolution-templeton-grant.page |website=University of Southampton|date=7 April 2016}}

Publications from the project include over 200 papers, a special issue,{{Cite web |title=Our special issue on Developmental Bias in Evolution is officially published online – Extended Evolutionary Synthesis |url=https://extendedevolutionarysynthesis.com/devobiasevo/ |access-date=2023-01-27 |language=en-GB}} and an anthology on Evolutionary Causation.{{Cite web |title=Evolutionary Causation |url=https://mitpress.mit.edu/9780262039925/evolutionary-causation/ |access-date=2023-01-27 |website=MIT Press |language=en-US}} In 2019 a final report of the 2016–2019 consortium was published, Putting the Extended Evolutionary Synthesis to the Test.[https://extendedevolutionarysynthesis.com/wp-content/uploads/2022/02/Final-Report-singlepage-compressed.pdf "Putting the Extended Evolutionary Synthesis to the Test]. extendedevolutionarysynthesis.com. Retrieved 22 August 2022.

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."[https://web.archive.org/web/20180702011327/https://evolution-institute.org/empowering-the-extended-evolutionary-synthesis/ "Empowering the Extended Evolutionary Synthesis"]. The Evolution Institute.

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.{{Cite journal|last=Craig|first=Lindsay R.|s2cid=84662773|date=June 2010|title=The So-Called Extended Synthesis and Population Genetics|journal=Biological Theory|volume=5|issue=2|pages=117–123|doi=10.1162/biot_a_00035|issn=1555-5542}} Proponents think that the conceptions of evolution at the core of the modern synthesis are too narrow{{cite journal |author=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 |title=Does Evolutionary Theory Need a Rethink? |journal=Nature |date=8 October 2014|volume=514|issue=7521 |pages=161–164 |doi=10.1038/514161a |pmid=25297418 |bibcode=2014Natur.514..161L|display-authors=etal|doi-access=free |hdl=1885/28950 |hdl-access=free }} 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.{{cite journal|last1=Noble|first1=Denis |author-link=Denis Noble |title=Evolution Beyond Neo-Darwinism: A New Conceptual Framework|journal=The Journal of Experimental Biology|date=1 January 2015|volume=218|issue=Pt 1|pages=7–13|doi=10.1242/jeb.106310|pmid=25568446 |doi-access=free}} 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.{{Cite journal |last=Müller |first=Gerd B. | author-link=Gerd B. Müller |date=2017-10-06|title=Why an extended evolutionary synthesis is necessary|journal=Interface Focus|volume=7|issue=5|pages=20170015|doi=10.1098/rsfs.2017.0015|issn=2042-8898|pmc=5566817|pmid=28839929}} In 2022, the John Templeton Foundation published a review of recent literature.{{Cite web |title=Extended Evolutionary Synthesis |url=https://www.templeton.org/discoveries/extended-evolutionary-synthesis |access-date=2023-01-27 |website=John Templeton Foundation}}

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