Extended evolutionary synthesis
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The extended evolutionary synthesis consists of a set of theoretical concepts 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. It includes multilevel selection, transgenerational epigenetic inheritance, niche construction, evolvability, and several concepts from evo-devo.
Not all biologists have agreed on the need for, or the scope of, an extended synthesis. 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"
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, one of the three founders of population genetics, and J. B. S. Haldane and Sewall Wright, between 1918 and 1932. Julian Huxley introduced the phrase "modern synthesis" in his 1942 book, Evolution: The Modern Synthesis.
During the 1950s, the English biologist C. H. Waddington called for an extended synthesis based on his research on epigenetics and genetic assimilation. An extended synthesis was also proposed by the Austrian zoologist Rupert Riedl, with the study of evolvability. In 1978, Michael J. D. White wrote about an extension of the modern synthesis based on new research from speciation.
1980s: 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. The ethologist John Endler wrote a paper in 1988 discussing processes of evolution that he felt had been neglected.
Contributions from 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 an 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. By contrast, a different strand of evo-devo following an organismal approach contributes to the extended synthesis by emphasizing (amongst others) developmental bias (both through facilitation and constraint), evolvability, and inherency of form  as primary factors in the evolution of complex structures and phenotypic novelties.
The idea of an extended synthesis was relaunched in 2007 by Massimo Pigliucci, 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. This includes:
- The role of prior configurations, genomic structures, and other traits in the organism in generating evolutionary variations.
- 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.
- The way that organismal development and developmental plasticity channel evolutionary pathways and generates phenotypic novelty
- How organisms modify the environments they belong to through niche construction.
Other processes such as evolvability, phenotypic plasticity, reticulate evolution, sex evolution and symbiogenesis are said by proponents to have been excluded or missed from the modern synthesis. 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. The biologist Eugene Koonin wrote in 2009 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."
The extended synthesis is characterized by its additional set of predictions that differ from the standard modern synthesis theory:
- change in phenotype can precede change in genotype
- changes in phenotype are predominantly positive, rather than neutral (see: neutral theory of molecular evolution)
- changes in phenotype are induced in many organisms, rather than one organism
- revolutionary change in phenotype can occur through mutation, facilitated variation or threshold events
- repeated evolution in isolated populations can be by convergent evolution or developmental bias
- 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)
- rapid evolution can result from simultaneous induction, natural selection and developmental dynamics
- biodiversity can be affected by features of developmental systems such as differences in evolvability
- heritable variation is directed towards variants that are adaptive and integrated with phenotype
- niche construction is biased towards environmental changes that suit the constructor's phenotype, or that of its descendants, and enhance their fitness
- kin selection
- multilevel selection
The extended evolutionary synthesis is currently being tested by a group of scientists from eight institutions in Britain, Sweden and the United States. The £7.7 million project is supported by a £5.7 million grant from the John Templeton Foundation.
The project is 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."
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 that the Extended synthesis is not radical enough. Proponents think that the conceptions of evolution at the core of the modern synthesis are too narrow. 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. 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.
- Wade, Michael J (2011). "The Neo-Modern Synthesis: The Confluence of New Data and Explanatory Concepts". BioScience. 61: 407–408. doi:10.1525/bio.2011.61.5.10.
- 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. ISSN 0962-8452. PMID 26246559.
- 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. doi:10.1038/nrg3028. PMID 21681209.
- Pigliucci, Massimo; Finkelman, Leonard (2014). "The Extended (Evolutionary) Synthesis Debate: Where Science Meets Philosophy". BioScience. 64: 511–516. doi:10.1093/biosci/biu062.
- 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. doi:10.1002/jez.b.22631.
- Müller, Gerd B. (December 2007). "Evo–devo: extending the evolutionary synthesis". Nature Reviews Genetics. 8 (12): 943–949. doi:10.1038/nrg2219. ISSN 1471-0056.
- 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. ISBN 0-309-06406-6. LCCN 99006259. OCLC 43803228.
The scientific consensus around evolution is overwhelming.
- Bock, Walter J. (July 1981). "Reviewed Work: The Evolutionary Synthesis. Perspectives on the Unification of Biology". The Auk. McLean, VA: American Ornithologists' Union. 98 (3): 644–646. ISSN 0004-8038. JSTOR 4086148.
- Fisher, Ronald A. (January 1919). "XV.—The Correlation between Relatives on the Supposition of Mendelian Inheritance". Transactions of the Royal Society of Edinburgh. London: Robert Grant & Son; Williams & Norgate. 52 (2): 399–433. doi:10.1017/S0080456800012163. ISSN 0080-4568. OCLC 4981124. "Paper read by J. Arthur Thomson on July 8, 1918 to the Royal Society of Edinburgh."
- 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 0-19-850440-3. LCCN 00702764. OCLC 45308589.
- Hubbs, C.L. (1943). "Evolution the new synthesis". American Naturalist. 77: 365–68. doi:10.1086/281134.
- Kimball, R.F. (1943). "The great biological generalization". Quarterly Review of Biology. 18: 364–67. doi:10.1086/394682.
- Karl P. Schmidt, Evolution the Modern Synthesis by Julian Huxley, Copeia, Vol. 1943, No. 4 (Dec. 31, 1943), pp. 262-263
- 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.
- Pigliucci, Massimo; et al. (2006). "Phenotypic plasticity and evolution by genetic assimilation". Journal of Experimental Biology. 209: 2362–2367. doi:10.1242/jeb.02070.CS1 maint: Explicit use of et al. (link)
- Huang S. (2011). The molecular and mathematical basis of Waddington’s epigenetic landscape: A framework for post-Darwinian biology? BioEssays 34: 149-157.
- 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.
- Parnell, Dennis R. (1978). Heralding a New Synthesis Modes of Speciation by M. J. D. White. Systematic Botany. Vol. 3, No. 1. p. 126.
- Gould, Stephen Jay. (1980). Is a New and General Theory of Evolution Emerging? Paleobiology. Vol. 6, No. 1. pp. 119-130.
- 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.
- "A More Modern Synthesis". American Scientist.
- 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.
- 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. DOI: 10.1146/annurev.es.19.110188.002143 JSTOR: 2097160
- 1937-2015,, Davidson, Eric H.,. The regulatory genome : gene regulatory networks in development and evolution. Amsterdam [Netherlands]. ISBN 0120885638. OCLC 61756485.
- Bateson, P (2005). "The Return of the Whole Organism". Journal of Biosciences. 30: 31–39. doi:10.1007/bf02705148. PMID 15824439.
- Huneman, Philippe (2010). "Assessing the Prospects for a Return of Organisms in Evolutionary Biology". History and Philosophy of the Life Sciences. 32: 341–372.
- Gilbert, S.F.; Opitz, G.; Raff, R. (1996). "Resynthesizing Evolutionary and Developmental Biology". Development and Evolution. 173: 357–372. doi:10.1006/dbio.1996.0032. PMID 8605997.
- Müller, G. B. (2007). "Evo-devo: Extending the evolutionary synthesis". Nature Reviews Genetics. 8: 943–949. doi:10.1038/nrg2219. PMID 17984972.
- "The Origins of Form". Natural History.
- 1941-, Hall, Brian K. (Brian Keith), (1998). Evolutionary developmental biology (2nd ed.). London: Chapman & Hall. ISBN 0412785803. OCLC 40606316.
- 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.
- Gerhart, John; Kirschner, Marc (2007-05-15). "The theory of facilitated variation". Proceedings of the National Academy of Sciences. 104 (suppl 1): 8582–8589. doi:10.1073/pnas.0701035104. PMC 1876433. PMID 17494755.
- 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.
- 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.
- Challenging the modern synthesis : adaptation, development, and inheritance. Huneman, Philippe,, Walsh, Denis M., 1958-. New York, NY. ISBN 9780199377183. OCLC 1001337947.
- 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, retrieved 2018-06-22
- 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, retrieved 2018-06-22
- 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
- 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.
- 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.
- Grant, Bob (1 January 2010). "Should Evolutionary Theory Evolve". The Scientist.
- Pigliucci, Massimo; Müller, Gerd B., eds. (26 March 2010). Evolution - the Extended Synthesis. The MIT Press. ISBN 978-0262513678.
- Meaden, Rhiannon (5 August 2015). "Redefining Evolutionary Biology". The Royal Society Publishing Blog.
- Indiana University (7 August 2015). "Expanding the Theory of Evolution". Lab Manager.
- 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.
- 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.
- 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.
- Moczek, Armin P. (2011-05-05). "Evolutionary biology: The origins of novelty". Nature. 473 (7345): 34–35. doi:10.1038/473034a. ISSN 0028-0836.
- 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. PMC 4632619. PMID 26246559.
- Lodé, Th (2011). "Sex is not a solution for reproduction, the libertine bubble theory". BioEssays. 33: 419–422. doi:10.1002/bies.201000125. PMID 21472739.
- Perez, JUlio E; Alfonsi, Carmen; Munoz, Carlos (2010). "Towards a New Evolutionary Theory" (PDF). Interciencia. 35: 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
- "Expanding Theory of Evolution". PhysOrg. 5 August 2015.
- Koonin, Eugene (2009). "Towards a postmodern synthesis of evolutionary biology". Cell Cycle. 8: 799–800. doi:10.4161/cc.8.6.8187. PMC 3410441. PMID 19242109.
- Lange, Axel; Nemeschkal, Hans L.; Müller, Gerd B. (April 2018). "A threshold model for polydactyly". Progress in Biophysics and Molecular Biology. doi:10.1016/j.pbiomolbio.2018.04.007. ISSN 0079-6107.
- 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.
- Laland, KN; Uller, T; Feldman, MW; Sterelny, K; Müller, GB; Moczek, A; Jablonka, E; Odling-Smee, J (Aug 2015). "The extended evolutionary synthesis: its structure, assumptions and predictions". Proc Biol Sci. 282: 20151019. doi:10.1098/rspb.2015.1019. PMC 4632619. PMID 26246559.
- Johnson BR, Lam SK (2010). Self-Organization, Natural Selection, and Evolution: Cellular Hardware and Genetic Software. Bioscience 60 (11): 879–885
- "Should Evolutionary Theory Evolve". University of Southampton. 7 April 2016.
- Laland, Kevin (21 December 2016). "Launching the Extended Evolutionary Synthesis project". Nature Research Ecology & Evolution Community.
- "Empowering the Extended Evolutionary Synthesis". The Evolution Institute.
- 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.
- 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.CS1 maint: Multiple names: authors list (link)
- 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. PMC 4436573. PMID 25568446.
- 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.
Defend the extended synthesis
- Arnold, Anthony J; Fristrup, Kurt (1982). "The Theory of Evolution by Natural Selection: A Hierarchical Expansion". Paleobiology. 8 (2): 113–129. doi:10.1017/s0094837300004462.
- Boto, Luis (2010). "Horizontal Gene Transfer in Evolution: Facts and Challenges". Proc Biol Sci. 277: 819–827. doi:10.1098/rspb.2009.1679. PMC 2842723. PMID 19864285.
- Carroll, Sean B (2008). "Evo-Devo and an Expanding Evolutionary Synthesis: A Genetic Theory of Morphological Evolution". Cell. 134: 25–36. doi:10.1016/j.cell.2008.06.030. PMID 18614008.
- Endler, John (1986). "The Newer Synthesis? Some Conceptual Problems in Evolutionary Biology". Oxford Surveys in Evolutionary Biology. 3: 224–243.
- Gilbert, Scott F. (2000). "A New Evolutionary Synthesis". In Developmental Biology, 6th edition. Sinauer. ISBN 0-87893-243-7
- Jablonka, Eva; Lamb, Marion J (2008). "Soft Inheritance: Challenging the Modern Synthesis". Genetics and Molecular Biology. 31: 389–395. doi:10.1590/s1415-47572008000300001.
- Koonin, Eugene (2009). "Towards a postmodern synthesis of evolutionary biology". Cell Cycle. 8: 799–800. doi:10.4161/cc.8.6.8187. PMC 3410441. PMID 19242109.
- Koonin, Eugene (2009). "The Origin at 150: is a new evolutionary synthesis in sight?". Trends Genet. 25: 473–475. doi:10.1016/j.tig.2009.09.007. PMC 2784144. PMID 19836100.
- Lodé, Thierry (2013). Manifeste pour une écologie évolutive, Darwin et après. Eds Odile Jacob, Paris.
- Müller, Gerd B (2014). "EvoDevo Shapes the Extended Synthesis". Biol Theory. 9: 119–121. doi:10.1007/s13752-014-0179-6.
- Pennisi, Elizabeth (2008). "Modernizing the Modern Synthesis". Science. 321: 196–197. doi:10.1126/science.321.5886.196.
- Postdarwinism: "The New Synthesis". A review of Ecological Developmental Biology: Integrating Epigenetics, Medicine, and Evolution, by Scott F. Gilbert and David Epel (Sinauer, 2009).
- "Post-modern synthesis?" A review of Developmental Plasticity and Evolution by Mary Jane West-Eberhard (Oxford University Press, 2003).
- Schrey; et al. (2012). "The Role of Epigenetics in Evolution: The Extended Synthesis". Genetics Research International. 2012: 286164.CS1 maint: Explicit use of et al. (link)
- Weber, Bruce H (2011). "Extending and Expanding the Darwinian Synthesis: The Role of Complex Systems Dynamics". Studies in History and Philosophy of Biological and Biomedical Science. 42: 75–81. doi:10.1016/j.shpsc.2010.11.014.
- Whitfield, John (2008). "Biology theory: Postmodern evolution?". Nature. 455: 281–284. doi:10.1038/455281a.
Criticism of the extended synthesis
- Coyne, Jerry (November 24, 2014). "Does evolution need a revolution?". Why Evolution Is True (Blog). Retrieved 2015-11-19.
- Craig, Lindsay (2010). "The So-Called Extended Synthesis and Population Genetics". Biological Theory. 5: 117–123. doi:10.1162/biot_a_00035.
- Dickens, Thomas; Rahman, Qazi. (2012). "The extended evolutionary synthesis and the role of soft inheritance in evolution". Proceedings of the Royal Society: B biological sciences, 279 (1740). pp. 2913–2921.
- Felsenstein, Joseph (1986). "Waiting for Post-Neo-Darwin". Evolution. 40 (4): 883–889. doi:10.2307/2408480.
- Haig, David (2007). "Weismann rules! OK? Epigenetics and the Lamarckian Temptation". Biology and Philosophy. 22: 415–428. doi:10.1007/s10539-006-9033-y.
- Kurland, CG; Canback, B; Berg, OG (2003). "Horizontal Gene Transfer: A Critical View". Proc Natl Acad Sci U S A. 100: 9658–9662. Bibcode:2003PNAS..100.9658K. doi:10.1073/pnas.1632870100. PMC 187805. PMID 12902542.
- Lynch, Michael (2007). "The frailty of adaptive hypotheses for the origins of organismal complexity". Proceedings of the National Academy of Sciences. 104: 8597–8604. Bibcode:2007PNAS..104.8597L. doi:10.1073/pnas.0702207104. PMC 1876435. PMID 17494740.
- Mayr, Ernst (2004). "80 years of watching the evolutionary scenery". Science. 305: 46–47. doi:10.1126/science.1100561.
- Merlin, Francesca (2010). "Evolutionary Chance Mutation: A Defense of the Modern Synthesis' Consensus View". Philosophy & Theory in Biology. 2: 22. doi:10.3998/ptb.6959004.0002.003.
- Stebbins, Ledyard G; Ayala, Francisco J (1981). "Is a New Evolutionary Synthesis Necessary?". Science. 213 (4511): 967–971. Bibcode:1981Sci...213..967L. doi:10.1126/science.213.4511.967.