Editing The Selfish Gene (section)

===Units of selection===
As to the [[unit of selection]]: "One internally consistent logical picture is that the unit of replication is the gene,...and the organism is one kind of ...entity on which selection acts directly."<ref name=Keller>{{cite book |title=Levels of Selection in Evolution |editor=Keller, Laurent |author1=Reeve, H. Kern |author2=Keller, Laurent  |chapter-url=https://books.google.com/books?id=5b_iEzNFlBUC&pg=PA5 |page=5 |chapter=Burying the debate over whether genes or individuals are the units of selection |isbn=978-0691007045 |year=1999 |publisher=Princeton University Press}}</ref> Dawkins proposed the matter without a distinction between 'unit of replication' and 'unit of selection' that he made elsewhere: "the fundamental unit of selection, and therefore of self-interest, is not the species, nor the group, nor even strictly the individual. It is the gene, the unit of heredity."<ref name=Dawkins76A>{{cite book |title=The Selfish Gene  |chapter=Chapter 1: Why are people? |chapter-url=https://books.google.com/books?id=WkHO9HI7koEC&pg=PA11 |page=11 |edition=30th Anniversary |isbn=978-0199291144 |year=2006 |publisher=Oxford University Press |author=Dawkins, Richard}}</ref> However, he continues in a later chapter:

:"On any sensible view of the matter Darwinian selection does not work on genes directly. ...The important differences between genes emerge only in their ''effects''. The technical word ''phenotype'' is used for the bodily manifestation of a gene, the effect that a gene has on the body...Natural selection favours some genes rather than others not because of the nature of the genes themselves, but because of their consequences—their phenotypic effects...But we shall now see that the phenotypic effects of a gene need to be thought of as ''all the effects that it has on the world''. ...The phenotypic effects of a gene are the tools by which it levers itself into the next generation. All I am going to add is that the tools may reach outside the individual body wall...Examples that spring to mind are artefacts like beaver dams, bird nests, and caddis houses." 
::— Richard Dawkins, ''The Selfish Gene'', Chapter 13, pp. 234, 235, 238

Dawkins's later formulation is in his book ''[[The Extended Phenotype]]'' (1982), where the process of selection is taken to involve every possible [[Phenotype|phenotypical]] effect of a gene.

[[Stephen Jay Gould]] finds Dawkins's position tries to have it both ways:<ref name=GouldSJ>{{cite book |title=The Structure of Evolutionary Theory |last=Gould |first=Stephen Jay |author-link=Stephen Jay Gould |url=https://books.google.com/books?id=6LMrAwAAQBAJ&pg=PA640 |pages=640–641 |isbn=978-0674417922 |year=2002 |publisher=Harvard University Press}}</ref>

:"Dawkins claims to prefer genes and to find greater insight in this formulation. But he allows that you or I might prefer organisms—and it really doesn't matter."
::— Stephen Jay Gould, ''The Structure of Evolutionary Theory'', pp. 640-641

The view of ''The Selfish Gene'' is that selection based upon groups and populations is rare compared to selection on individuals. Although supported by Dawkins and by many others, this claim continues to be disputed.<ref name=Pinker>For a view opposing 'multilevel selection', see {{cite web |author=Pinker, Steven |author-link=Steven Pinker |title=The false allure of group selection |url=http://edge.org/conversation/the-false-allure-of-group-selection |work=Edge: the online version of "The Reality Club" |publisher=John Brockman |date=18 June 2012 |access-date=1 March 2015}} For a more modern (and technical) argument, see {{cite journal |title=Kin selection is the key to altruism |author1=Foster, Kevin R. |author2=Wenseleers, Tom |author3=Francis L.W. Ratnieks |url=https://lirias.kuleuven.be/bitstream/123456789/69591/1/foster_etal_tree_2006_kin_selection_key_to_altruism.pdf |archive-url=https://web.archive.org/web/20150402131630/https://lirias.kuleuven.be/bitstream/123456789/69591/1/foster_etal_tree_2006_kin_selection_key_to_altruism.pdf |archive-date=2015-04-02 |url-status=live |journal=Trends in Ecology & Evolution |volume=21 |pages=57–60 |number=2 |date=February 2006 |doi=10.1016/j.tree.2005.11.020|pmid=16701471 |bibcode=2006TEcoE..21...57F }}</ref><ref name=Smaldino>For a view supporting 'evolutionary competition between cultural groups', see {{cite journal |author=Smaldino, Paul E. |title=The cultural evolution of emergent group-level traits |journal=Behavioral and Brain Sciences |volume= 37 |issue=3 |year=2014 |pages=243–254 |url=http://www.zoology.ubc.ca/~doebeli/reprints/smaldino2014.pdf |archive-url=https://web.archive.org/web/20150402094752/http://www.zoology.ubc.ca/~doebeli/reprints/smaldino2014.pdf |archive-date=2015-04-02 |url-status=live |doi=10.1017/s0140525x13001544|pmid=24970399 |s2cid=52872815 }}</ref> While naïve versions of [[group selectionism]] have been disproved, more sophisticated formulations make accurate predictions in some cases while positing selection at higher levels.<ref>{{cite journal |last=Wilson |first=David Sloan |author2=Wilson, Edward O.  |year=2007 |title=Rethinking the Theoretical Foundations of Sociobiology |journal=[[The Quarterly Review of Biology]] |jstor=10.1086/522809|volume=82 |issue=4 |pages=327–348 |doi=10.1086/522809 |pmid=18217526|s2cid=37774648 }}</ref> Both sides agree that very favourable genes are likely to prosper and replicate if they arise and both sides agree that living in groups can be an advantage to the group members. The conflict arises in part over defining concepts: 
:"Cultural evolutionary theory, however, has suffered from an overemphasis on the experiences and behaviors of individuals at the expense of acknowledging complex group organization...Many important behaviors related to the success and function of human societies are only properly defined at the level of groups".<ref name=Smaldino/>

In ''[[The Social Conquest of Earth]]'' (2012), the entomologist [[E. O. Wilson]] contends that although the selfish-gene approach was accepted "until 2010 [when] [[Martin Nowak]], [[Corina Tarnita]], and I demonstrated that inclusive fitness theory, often called kin selection theory, is both mathematically and biologically incorrect."<ref name=Wilson>{{cite book |last=Wilson |first=Edward |title=The Social Conquest of Earth |year=2012 |publisher=Liveright Publishing |location=New York, NY |isbn=978-0-87140-413-8 |page=[https://archive.org/details/socialconquestof0000wils/page/143 143] |url=https://archive.org/details/socialconquestof0000wils|url-access=registration }} Although it contains no reference to the "selfish gene", Wilson probably is referring to {{cite journal |title=The evolution of eusociality |author1=Nowak, M. A. |author2=Tarnita, C. E. |author3=Wilson, Edward O. |name-list-style=amp |journal=Nature |date=26 August 2010 |pages=1057–1062 |doi=10.1038/nature09205 |pmc=3279739 |pmid=20740005 |volume=466|issue=7310 |bibcode=2010Natur.466.1057N }} and {{cite journal |title=Natural selection versus kin selection | pmc=3279739 | pmid=20740005 |doi=10.1038/nature09205 |volume=466 |issue=7310 |date=August 2010 |journal=Nature |pages=1057–62 | last1 = Nowak | first1 = MA | last2 = Tarnita | first2 = CE | last3 = Wilson | first3 = EO| bibcode=2010Natur.466.1057N }}; Supplementary Information for ''The evolution of eusociality''</ref> Chapter 18 of ''The Social Conquest of Earth'' describes the deficiencies of kin selection and outlines group selection, which Wilson argues is a more realistic model of social evolution. He criticises earlier approaches to social evolution, saying: "unwarranted faith in the central role of kinship in social evolution has led to the reversal of the usual order in which biological research is conducted. The proven best way in evolutionary biology, as in most of science, is to define a problem arising during empirical research, then select or devise the theory that is needed to solve it. Almost all research in inclusive-fitness theory has been the opposite:  hypothesize the key roles of kinship and kin selection, then look for evidence to test that hypothesis." According to Wilson: "People must have a tribe...Experiments conducted over many years by social psychologists have revealed how swiftly and decisively people divide into groups, and then discriminate in favor of the one to which they belong." (pp.&nbsp;57, 59) According to Wilson: "Different parts of the brain have evolved by group selection to create groupishness." (p.&nbsp;61)

Some authors consider facets of this debate between Dawkins and his critics about the level of selection to be blather:<ref name=Keller2>{{cite book |title=Levels of Selection in Evolution |author1=H Kern Reeve |author2=Laurent Keller |editor=Laurent Keller |chapter-url=https://books.google.com/books?id=5b_iEzNFlBUC&pg=PA4 |page=4 |chapter=Burying the debate over whether genes or individuals are the units of selection |isbn=978-0691007045 |year=1999 |publisher=Princeton University Press}}</ref>
:"The particularly frustrating aspects of these constantly renewed debates is that, even though they seemed to be sparked by rival theories about how evolution works, in fact they often involve only rival metaphors for the very same evolutionary logic and [the debates over these aspects] are thus empirically empty."
::— Laurent Keller, ''Levels of Selection in Evolution'', p.4

Other authors say Dawkins has failed to make some critical distinctions, in particular, the difference between group selection for group advantage and group selection conveying individual advantage.<ref name=Lloyd>{{cite book |title=The Cambridge Companion to the Philosophy of Biology |author=Lloyd, Elizabeth |chapter=Units and Levels of Selection |pages=54–57 |isbn=978-1139827621 |year=2007 |publisher=Cambridge University Press |chapter-url=https://books.google.com/books?id=aZOgg-x4UyIC&pg=PA54 |editor1=Hull, David L. |editor2=Ruse, Michael}}</ref>