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===The birth of evo-devo and a second synthesis=== In 1977, a revolution in thinking about evolution and developmental biology began, with the arrival of [[recombinant DNA]] technology in [[genetics]], the book ''Ontogeny and Phylogeny'' by [[Stephen J. Gould]] and the paper [[Evolutionary tinkering|"Evolution and Tinkering"]]<ref name="Jacob 1977">{{Cite journal |last=Jacob |first=François |date=10 June 1977 |title=Evolution and Tinkering |journal=Science |volume=196 |issue=4295 |pages=1161–1166 |bibcode=1977Sci...196.1161J |doi=10.1126/science.860134 |pmid=860134}}</ref> by [[François Jacob]]. Gould laid to rest Haeckel's interpretation of evolutionary embryology, while Jacob set out an alternative theory.<ref name=Gilbert2003/> This led to [[extended evolutionary synthesis|a second synthesis]],<ref>{{Cite journal |last=Gilbert |first=S.F. |last2=Opitz |first2=J.M. |last3=Raff |first3=R.A. |date=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}}</ref><ref>{{Cite journal |last=Müller |first=G. B. |author-link=Gerd B. Müller |date=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 |s2cid=19264907}}</ref> at last including embryology as well as [[molecular genetics]], phylogeny, and evolutionary biology to form evo-devo.<ref>{{Cite journal |last=Goodman |first=C. S. |last2=Coughlin |first2=B. C. |year=2000 |editor2-last=Coughlin B. S. |title=Special feature: The evolution of evo-devo biology |journal=[[Proceedings of the National Academy of Sciences]] |volume=97 |issue=9 |pages=4424–4456 |bibcode=2000PNAS...97.4424G |doi=10.1073/pnas.97.9.4424 |pmc=18255 |pmid=10781035 |doi-access=free |editor1=Goodman, C. S.}}</ref><ref>{{Cite journal |last=[[Gerd Müller (theoretical biologist)|Müller GB]] and [[Stuart Newman|Newman SA]] (Eds.) |year=2005 |title=Special issue: Evolutionary Innovation and Morphological Novelty |url=http://www3.interscience.wiley.com/cgi-bin/jissue/112149101 |journal=Journal of Experimental Zoology Part B |volume=304B |issue=6 |pages=485–631 |doi=10.1002/jez.b.21080 |pmid=16252267 |archive-url=https://archive.today/20121211055145/http://www3.interscience.wiley.com/cgi-bin/jissue/112149101 |archive-date=2012-12-11}}</ref> In 1978, [[Edward B. Lewis]] discovered [[homeosis|homeotic]] genes that regulate embryonic development in ''[[Drosophila]]'' fruit flies, which like all insects are [[arthropods]], one of the major [[Phylum|phyla]] of invertebrate animals.<ref name="Palmer 2004">{{Cite journal |last=Palmer |first=R.A. |year=2004 |title=Symmetry breaking and the evolution of development |journal=[[Science (journal)|Science]] |volume=306 |issue=5697 |pages=828–833 |bibcode=2004Sci...306..828P |citeseerx=10.1.1.631.4256 |doi=10.1126/science.1103707 |pmid=15514148 |s2cid=32054147}}</ref> [[Bill McGinnis]] quickly discovered homeotic gene sequences, [[homeobox]]es, in animals in other phyla, in [[vertebrate]]s such as [[frog]]s, [[bird]]s, and [[mammals]]; they were later also found in [[fungi]] such as [[yeast]]s, and in [[plant]]s.<ref name=Winchester/><ref>{{Cite web |last=Bürglin |first=Thomas R. |title=The Homeobox Page |url=http://homeobox.biosci.ki.se/ |access-date=13 October 2016 |publisher=[[Karolinska Institutet]]}}</ref> There were evidently strong similarities in the genes that controlled development across all the [[eukaryote]]s.<ref>{{Cite journal |last=Holland |first=P.W. |date=2013 |title=Evolution of homeobox genes |journal=Wiley Interdiscip Rev Dev Biol |volume=2 |issue=1 |pages=31–45 |doi=10.1002/wdev.78 |pmid=23799629 |s2cid=44396110 |quote=Homeobox genes are found in almost all eukaryotes, and have diversified into 11 gene classes and over 100 gene families in animal evolution, and 10 to 14 gene classes in plants.}}</ref> In 1980, [[Christiane Nüsslein-Volhard]] and [[Eric Wieschaus]] described [[gap gene]]s which help to create the segmentation pattern in [[Drosophila embryogenesis|fruit fly embryos]];<ref name="Nusslein">{{Cite journal |last=Nüsslein-Volhard, C. |last2=Wieschaus, E. |date=October 1980 |title=Mutations affecting segment number and polarity in ''Drosophila'' |journal=Nature |volume=287 |issue=5785 |pages=795–801 |bibcode=1980Natur.287..795N |doi=10.1038/287795a0 |pmid=6776413 |s2cid=4337658}}</ref><ref name="Arthur2002">{{Cite journal |last=Arthur |first=Wallace |date=14 February 2002 |title=The emerging conceptual framework of evolutionary developmental biology |journal=Nature |volume=415 |issue=6873 |pages=757–764 |bibcode=2002Natur.415..757A |doi=10.1038/415757a |pmid=11845200 |s2cid=4432164}}</ref> they and Lewis won a [[Nobel Prize in Physiology or Medicine|Nobel Prize]] for their work in 1995.<ref name="Winchester">{{Cite journal |last=Winchester |first=Guil |year=2004 |title=Edward B. Lewis 1918-2004 |url=http://www.cell.com/current-biology/pdf/S0960-9822(04)00683-9.pdf |publication-date=Sep 21, 2004 |volume=14 |issue=18 |pages=R740–742 |doi=10.1016/j.cub.2004.09.007 |pmid=15380080 |s2cid=32648995 |doi-access=free |periodical=Current Biology}}</ref><ref>{{Cite web |title=Eric Wieschaus and Christiane Nüsslein-Volhard: Collaborating to Find Developmental Genes |url=https://www.ibiology.org/ibiomagazine/eric-wieschaus-and-christiane-nusselin-volhard.html |url-status=dead |archive-url=https://web.archive.org/web/20161013223611/https://www.ibiology.org/ibiomagazine/eric-wieschaus-and-christiane-nusselin-volhard.html |archive-date=13 October 2016 |access-date=13 October 2016 |publisher=iBiology}}</ref> Later, more specific similarities were discovered: for example, the [[distal-less]] gene was found in 1989 to be involved in the development of appendages or limbs in fruit flies,<ref>{{Cite journal |last=Cohen, S. M. |last2=Jurgens, G. |date=1989 |title=Proximal-distal pattern formation in Drosophila: cell autonomous requirement for Distal-less activity in limb development |journal=EMBO J. |volume=8 |issue=7 |pages=2045–2055 |doi=10.1002/j.1460-2075.1989.tb03613.x |pmc=401088 |pmid=16453891}}</ref> the fins of fish, the wings of chickens, the [[parapodia]] of marine [[annelid]] worms, the ampullae and siphons of tunicates, and the [[tube feet]] of [[sea urchin]]s. It was evident that the gene must be ancient, dating back to the [[Urbilaterian|last common ancestor of bilateral animals]] (before the [[Ediacaran]] Period, which began some 635 million years ago). Evo-devo had started to uncover the ways that all animal bodies were built during development.<ref>{{Cite book |last=Carroll |first=Sean B. |author-link=Sean B. Carroll |title=Endless Forms Most Beautiful: The New Science of Evo Devo and the Making of the Animal Kingdom |title-link=Endless Forms Most Beautiful (book) |date=2006 |publisher=Weidenfeld & Nicolson [Norton] |isbn=978-0-297-85094-6 |pages=63–70 |orig-date=2005}}</ref><ref>{{Cite journal |last=Panganiban |first=G. |last2=Irvine |first2=S. M. |last3=Lowe |first3=C. |last4=Roehl |first4=H. |last5=Corley |first5=L. S. |last6=Sherbon |first6=B. |last7=Grenier |first7=J. K. |last8=Fallon |first8=J. F. |last9=Kimble |first9=J. |last10=Walker |first10=M. |last11=Wray |first11=G. A. |last12=Swalla |first12=B. J. |last13=Martindale |first13=M. Q. |last14=Carroll |first14=S. B. |year=1997 |title=The origin and evolution of animal appendages |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=94 |issue=10 |pages=5162–5166 |bibcode=1997PNAS...94.5162P |doi=10.1073/pnas.94.10.5162 |pmc=24649 |pmid=9144208 |doi-access=free}}</ref>
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