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===Selectionism and sociobiology=== Gould was a champion of [[biological constraints]], internal limitations upon developmental pathways, as well as other non-selectionist forces in evolution. Rather than direct [[adaptation]]s, he considered many higher functions of the [[human brain]] to be the unintended [[Unintended consequence|side consequence]] of [[natural selection]].<ref name="Spandrels"/> To describe such co-opted features, he coined the term [[exaptation]] with paleontologist [[Elisabeth Vrba]].<ref>{{Citation | last1 = Gould | first1 = S. J. | last2 = Vrba | first2 = E. | year = 1982 | title = Exaptation—a missing term in the science of form | url = http://www2.hawaii.edu/~khayes/Journal_Club/fall2006/Gould_&_Vrb_1982_Paleobio.pdf | journal = Paleobiology | volume = 8 | issue = 1| pages = 4–15 | postscript = . | doi = 10.1017/S0094837300004310 | bibcode = 1982Pbio....8....4G | s2cid = 86436132 }}</ref> Gould believed this feature of human mentality undermines an [[Genetic determinism|essential premise]] of human [[sociobiology]] and [[evolutionary psychology]].<ref>{{cite web |url=https://embryo.asu.edu/pages/spandrels-san-marco-and-panglossian-paradigm-critique-adaptationist-programme-1979-stephen-j |title="The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme" (1979), by Stephen J. Gould and Richard C. Lewontin | Embryo Project Encyclopedia }}</ref><ref>{{cite web |url=https://www.livescience.com/39688-exaptation.html |title=Exaptation: How Evolution Uses What's Available |website=[[Live Science]] |date=September 16, 2013 }}</ref> ====Against ''Sociobiology''==== In 1975, Gould's Harvard colleague [[E. O. Wilson]] introduced his analysis of animal behavior (including human behavior) based on a sociobiological framework that suggested that many social behaviors have a strong evolutionary basis.<ref> Wilson, E. O. (1975). ''[[Sociobiology: The New Synthesis]]''. Cambridge MA: Harvard University Press.</ref> In response, Gould, [[Richard Lewontin]], and [[Sociobiology Study Group|others]] from the Boston area wrote the subsequently well-referenced letter to ''[[The New York Review of Books]]'' entitled, "Against 'Sociobiology'". This [[open letter]] criticized Wilson's notion of a "deterministic view of human society and human action".<ref> Allen, Elizabeth, et al. (1975). [http://www.nybooks.com/articles/9017?sess=305fe41afae729849e1e7eb4b004bb81 "Against 'Sociobiology'".] [letter] ''[[New York Review of Books]]'' 22 (Nov. 13): 182, 184–186.</ref> But Gould did not rule out sociobiological explanations for many aspects of animal behavior, and later wrote: "Sociobiologists have broadened their range of selective stories by invoking concepts of [[inclusive fitness]] and [[kin selection]] to solve (successfully I think) the vexatious problem of [[altruism]]—previously the greatest stumbling block to a Darwinian theory of social behavior... Here sociobiology has had and will continue to have success. And here I wish it well. For it represents an extension of basic Darwinism to a realm where it should apply."<ref>Gould, S. J. (1980). "Sociobiology and the Theory of Natural Selection". In G. W. Barlow and J. Silverberg, eds., ''Sociobiology: Beyond Nature/Nurture?'' Boulder CO: Westview Press, pp. 257–269.</ref> {{anchor|Panglossian Paradigm|panglossian paradigm|Panglossian paradigm}} ====Spandrels and the Panglossian paradigm==== {{further|Spandrel (biology)}} [[Image:Kostel Nejsvětější Trojice (Fulnek) – frs-002.jpg|thumb|left|A spandrel from the [[Most Holy Trinity Church, Fulnek|Holy Trinity Church]] in Fulnek, [[Czech Republic]].]] With Richard Lewontin, Gould wrote an influential 1979 paper entitled, "[[The Spandrels of San Marco and the Panglossian Paradigm]]",<ref name="Spandrels">{{cite journal | last1 = Gould | first1 = S. J. | last2 = Lewontin | first2 = Richard | year = 1979 | title = The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme | journal = Proc. R. Soc. Lond. B Biol. Sci. | volume = 205 | issue = 1161| pages = 581–98 | doi = 10.1098/rspb.1979.0086 | pmid = 42062 | bibcode = 1979RSPSB.205..581G | s2cid = 2129408 }} for background see Gould's [http://www.edge.org/documents/ThirdCulture/i-Ch.2.html "The Pattern of Life's History"] {{Webarchive|url=https://web.archive.org/web/20150414021048/http://www.edge.org/documents/ThirdCulture/i-Ch.2.html |date=April 14, 2015 }} in John Brockman ''[http://www.edge.org/documents/ThirdCulture/d-Contents.html The Third Culture] {{Webarchive|url=https://web.archive.org/web/20160130235501/http://www.edge.org/documents/ThirdCulture/d-Contents.html |date=January 30, 2016 }}''. New York: Simon & Schuster. 1996, pp. 52–64. {{ISBN|0-684-82344-6}}.</ref> which introduced the architectural term "[[spandrel]]" into evolutionary biology. In architecture, a spandrel is a triangular space which exists over the [[Haunch (arch)|haunch]]es of an arch.<ref>ITC (1908) ''International Library of Technology'' 38 (3): [https://books.google.com/books?id=Yp9IAAAAMAAJ&pg=RA7-PA22 22.]</ref><ref name="MarkSpandrel">Mark, Robert (1996). [https://www.uv.mx/personal/tcarmona/files/2010/08/Mark-1996.pdf "Architecture and Evolution"] ''American Scientist'' (July–August): 383-389.</ref> Spandrels—more often called [[pendentive]]s in this context—are found particularly in classical architecture, especially Byzantine and Renaissance churches. When visiting [[Venice]] in 1978, Gould noted that the spandrels of the [[St Mark's Basilica|San Marco]] cathedral, while quite beautiful, were not spaces planned by the architect. Rather the spaces arise as "necessary architectural byproducts of mounting a dome on rounded arches." Gould and Lewontin thus defined "[[spandrel (biology)|spandrels]]" in the evolutionary biology context to mean any biological feature of an organism that arises as a necessary side consequence of other features, which is not directly selected for by natural selection. Proposed examples include the "masculinized genitalia in female [[Hyenas#Appearance and biology|hyenas]], exaptive use of an [[Umbilicus (mollusc)|umbilicus]] as a brooding chamber by snails, the shoulder hump of the giant [[Irish Elk|Irish deer]], and several key features of human mentality".<ref>{{Cite journal |doi=10.1073/pnas.94.20.10750 |pmid=11038582|pmc=23474 |title=The exaptive excellence of spandrels as a term and prototype |journal=Proceedings of the National Academy of Sciences |volume=94 |issue=20 |pages=10750–10755 |year=1997 |last=Gould |first=S. J. |bibcode=1997PNAS...9410750G |doi-access=free}}</ref> In [[Voltaire]]'s ''[[Candide]]'', [[Candide#Synopsis|Dr. Pangloss]] is portrayed as a clueless scholar who, despite the evidence, insists that "all is for the best in this best of all possible worlds". Gould and Lewontin asserted that it is [[Panglossian]] for evolutionary biologists to view all traits as atomized things that had been naturally selected for, and criticised biologists for not granting theoretical space to other causes, such as phyletic and developmental [[Biological constraints|constraints]]. The relative frequency of spandrels, so defined, versus adaptive features in nature, remains a controversial topic in [[evolutionary biology]].<ref>{{cite news |last=Maynard Smith |first=John |date=November 30, 1995 |title=Genes, Memes, & Minds |url=http://www.nybooks.com/articles/1703 |newspaper=[[The New York Review of Books]] |pages=46–48 |quote=By and large, I think their [Spandrels] paper had a healthy effect. ... Their critique forced us to clean up our act and to provide evidence for our stories. But adaptationism remains the core of biological thinking.}}</ref><ref>{{cite journal|first=Ernst|last=Mayr|author-link=Ernst Mayr|url=http://academic.reed.edu/biology/courses/bio342/2010_syllabus/2010_readings/Mayr_1983.pdf|title=How to Carry Out the Adaptationist Program?|journal=[[The American Naturalist]]|date=March 1983|volume=121|issue=3|pages=324–334|doi=10.1086/284064|bibcode=1983ANat..121..324M |s2cid=3937726}}</ref><ref>{{cite book|first=George C.|last=Williams|author-link=George C. Williams (biologist)|title=Natural Selection: Domains, Levels, and Challenges|publisher=[[Oxford University Press]]|location=New York City|date=1992|isbn=978-0195069334}}</ref> An illustrative example of Gould's approach can be found in [[Elisabeth Lloyd]]'s case study suggesting that the female orgasm is a by-product of shared developmental pathways.<ref name="caseofthefemaleorgasm">{{cite book |last=Lloyd |first=Elisabeth Anne |title=The Case of The Female Orgasm: Bias in the science of evolution |publisher=Harvard University Press |publication-place=Cambridge, Mass. |year=2005 |isbn=978-0-674-04030-4 |oclc=432675780 |page={{page needed|date=June 2022}} |url=https://archive.org/details/caseoffemaleorga0000lloy |url-access=registration |via=Internet Archive}}</ref> Gould also wrote on this topic in his essay "Male Nipples and Clitoral Ripples," prompted by Lloyd's earlier work.<ref name="MaleNipples">Gould, S.J. (1992). [https://www.pdf-archive.com/2018/03/26/sjgould-male-nipples-and-clitoral-ripples/sjgould-male-nipples-and-clitoral-ripples.pdf "Male Nipples and Clitoral Ripples".] {{Webarchive|url=https://web.archive.org/web/20190922040512/https://www.pdf-archive.com/2018/03/26/sjgould-male-nipples-and-clitoral-ripples/sjgould-male-nipples-and-clitoral-ripples.pdf |date=September 22, 2019 }} In ''[[Bully for Brontosaurus]]: Further Reflections in Natural History''. New York: W. W. Norton. pp. 124–138.</ref> Gould was criticized by philosopher [[Daniel Dennett]] for using the term spandrel instead of pendentive,<ref>Dennett, Daniel (1995) ''Darwin's Dangerous Idea''. New York: Penguin Books, [https://books.google.com/books?id=Y77BAwAAQBAJ&pg=PA274 p. 272.]</ref> a spandrel that curves across a right angle to support a dome. Robert Mark, a professor of civil engineering at Princeton, offered his expertise in the pages of ''[[American Scientist]]'', noting that these definitions are often misunderstood in [[architectural theory]]. Mark concluded, "Gould and Lewontin's misapplication of the term spandrel for pendentive perhaps implies a wider latitude of design choice than they intended for their analogy. But Dennett's critique of the architectural basis of the analogy goes even further astray because he slights the technical rationale of the architectural elements in question."<ref name="MarkSpandrel"/>
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