Contents 1 Good of the species 2 Selfish selection 3 Group selection revisited 3.1 Multilevel selection theory 3.2 Other arguments 3.3 Gene-culture coevolution in humans 3.4 Differing evolutionarily stable strategies (ESSs) 3.5 Implications in population biology 4 Criticism 5 References 6 Further reading 7 External links

Good of the species[edit] Once Darwinism had been accepted, animal behavior was glibly explained with unsubstantiated hypotheses about survival value, which was largely taken for granted. The naturalist Konrad Lorenz had argued loosely in books like On Aggression (1966) that animal behavior patterns were "for the good of the species",[1][2] without actually studying survival value in the field; the ethologist Niko Tinbergen praised Lorenz for his interest in the survival value of behavior, and naturalists enjoyed Lorenz's writings for the same reason.[2] In 1962, group selection was used as a popular explanation for adaptation by the zoologist V. C. Wynne-Edwards.[3][4]

Selfish selection[edit] Social behavior in honeybees is explained by kin selection: their haplodiploid inheritance system makes workers very closely related to their queen (centre). Further information: natural selection and kin selection From the mid 1960s, evolutionary biologists argued that natural selection acted primarily at the level of the individual. In 1964, John Maynard Smith,[5] C.M. Perrins (1964),[6] and George C. Williams in his 1966 book Adaptation and Natural Selection cast serious doubt on group selection as a major mechanism of evolution; Williams's 1971 book Group Selection assembled writings from many authors on the same theme.[7][8] The great majority of evolutionary biologists believe (2011) that selection above the level of the individual is a special case, probably limited to the unique inheritance system (involving haplodiploidy) of the eusocial Hymenoptera such as honeybees, which encourages kin selection.[9]

Group selection revisited[edit] Further information: Unit of selection Experiments from the late 1970s suggested that group selection involving altruism was possible.[10] Early group selection models were flawed because they assumed that genes acted independently; but genetically-based interactions among individuals are ubiquitous in group formation because genes must cooperate for the benefit of association in groups to enhance the fitness of group members.[11] Additionally, group selection on the level of the species is flawed because it is difficult to see how selective pressures would be applied; selection in social species of groups against other groups, rather than the species entire, seems to be the level at which selective pressures are plausible. On the other hand, kin selection is accepted as an explanation of altruistic behavior.[12] Since the 1990s, group selection models have seen a resurgence.[13][14][15][16][17] Some biologists argue that kin selection and multilevel selection are both needed to "obtain a complete understanding of the evolution of a social behavior system".[18] In 2010 three authors including E. O. Wilson argued for multi-level selection, including group selection, to correct what they saw as deficits in the explanatory power of inclusive fitness.[19] The response was a back-lash from 137 other evolutionary biologists who argued "that their arguments are based upon a misunderstanding of evolutionary theory and a misrepresentation of the empirical literature".[9] Multilevel selection theory[edit] In 1994 David Sloan Wilson and Elliott Sober argued that the case against group selection had been overstated. They considered whether groups can have functional organization in the same way as individuals, and consequently whether groups can be "vehicles" for selection. They do not posit evolution on the level of the species, but selective pressures that winnow out small groups within a species, e.g. groups of social insects or primates. Groups that cooperate better might survive and reproduce more than those that did not. Resurrected in this way, Wilson & Sober's new group selection is called multilevel selection theory.[20] David Sloan Wilson and Elliott Sober's 1994 Multilevel Selection Model, illustrated by a nested set of Russian matryoshka dolls. Wilson himself compared his model to such a set. Wilson compared the layers of competition and evolution to nested sets of Russian matryoshka dolls.[15] The lowest level is the genes, next come the cells, then the organism level and finally the groups. The different levels function cohesively to maximize fitness, or reproductive success. The theory asserts that selection for the group level, involving competition between groups, must outweigh the individual level, involving individuals competing within a group, for a group-beneficiating trait to spread.[21] Multilevel selection theory focuses on the phenotype because it looks at the levels that selection directly acts upon.[15] For humans, social norms can be argued to reduce individual level variation and competition, thus shifting selection to the group level. Wilson ties the multilevel selection theory regarding humans to another theory, gene-culture coevolution, by acknowledging that culture seems to characterize a group-level mechanism for human groups to adapt to environmental changes.[21] MLS theory can be used to evaluate the balance between group selection and individual selection in specific cases.[21] An experiment by William Muir compared egg productivity in hens, showing that a hyper-aggressive strain had been produced through individual selection, leading to many fatal attacks after only six generations; by implication, it could be argued that group selection must have been acting to prevent this in real life.[22] Group selection has most often been postulated in humans and, notably, eusocial Hymenoptera that make cooperation a driving force of their adaptations over time and have a unique system of inheritance involving haplodiploidy that allows the colony to function as an individual while only the queen reproduces.[23] Wilson & Sober's work revived interest in multilevel selection. In a 2005 article,[24] E. O. Wilson argued that kin selection could no longer be thought of as underlying the evolution of extreme sociality, for two reasons. First, he suggested, the argument that haplodiploid inheritance (as in the Hymenoptera) creates a strong selection pressure towards nonreproductive castes is mathematically flawed.[25] Second, eusociality no longer seems to be confined to the hymenopterans; increasing numbers of highly social taxa have been found in the years since Wilson's foundational text on sociobiology was published in 1975,[26] including a variety of insect species, as well as two rodent species (the naked mole-rat and the Damaraland mole rat). Wilson suggests the equation for Hamilton's rule:[27] rb > c (where b represents the benefit to the recipient of altruism, c the cost to the altruist, and r their degree of relatedness) should be replaced by the more general equation rbk + be > c in which bk is the benefit to kin (b in the original equation) and be is the benefit accruing to the group as a whole. He then argues that, in the present state of the evidence in relation to social insects, it appears that be>rbk, so that altruism needs to be explained in terms of selection at the colony level rather than at the kin level. However, kin selection and group selection are not distinct processes, and the effects of multi-level selection are already accounted for in Hamilton's rule, rb>c,[28] provided that an expanded definition of r, not requiring Hamilton's original assumption of direct genealogical relatedness, is used, as proposed by E. O. Wilson himself.[29] Other arguments[edit] Spatial populations of predators and prey show restraint of reproduction at equilibrium, both individually[30] and through social communication,[31] as originally proposed by Wynne-Edwards. While these spatial populations do not have well-defined groups for group selection, the local spatial interactions of organisms in transient groups are sufficient to lead to a kind of multi-level selection. There is however as yet no evidence that these processes operate in the situations where Wynne-Edwards posited them; Rauch et al.'s analysis,[30] for example, is of a host-parasite situation, which was recognised as one where group selection was possible even by E. O. Wilson (1975), in a treatise broadly hostile to the whole idea of group selection.[26] Specifically, the parasites do not individually moderate their transmission; rather, more transmissible variants "continually arise and grow rapidly for many generations but eventually go extinct before dominating the system." A variant of group selection theory rooted in the idea of population viscosity (limited offspring dispersal), first proposed by Hamilton (1964), is widely present in natural populations. This population structure builds a continuum between individual selection, kin selection, kin group selection and group selection without a clear boundary for each level. However, early theoretical models by D.S. Wilson et al. (1992)[32] and Taylor (1992)[33] showed that pure population viscosity cannot lead to cooperation and altruism. This is because any benefit generated by kin cooperation is exactly cancelled out by kin competition; additional offspring from cooperation are eliminated by local competition. However, this exact cancelling out also suggests that any additional benefit of local cooperation would be sufficient to allow cooperation to evolve.[16] Mitteldorf and D.S. Wilson (2000) later showed that if the population is allowed to fluctuate, then local populations can temporarily store the benefit of local cooperation and promote the evolution of cooperation and altruism.[34] By assuming individual differences in adaptations, Yang (2013) further showed that the benefit of local altruism can be stored in the form of offspring quality and thus promote the evolution of altruism even if the population does not fluctuate. This is because local competition among more individuals resulting from local altruism increases the average local fitness of the individuals that survive.[16] Gene-culture coevolution in humans[edit] Main articles: Dual inheritance theory and cultural group selection Humanity has developed extremely rapidly, arguably through gene-culture coevolution, leading to complex cultural artefacts like the gopuram of the Sri Mariammam temple, Singapore. Gene-culture coevolution (also called dual inheritance theory) is a modern hypothesis (applicable mostly to humans) that combines evolutionary biology and modern sociobiology to indicate group selection.[35] It treats culture as a separate evolutionary system that acts in parallel to the usual genetic evolution to transform human traits. It is believed that this approach of combining genetic influence with cultural influence over several generations is not present in the other hypotheses such as reciprocal altruism and kin selection, making gene-culture evolution one of the strongest realistic hypotheses for group selection. Fehr provides evidence of group selection taking place in humans presently with experimentation through logic games such as prisoner’s dilemma, the type of thinking that humans have developed many generations ago.[36] Gene-culture coevolution allows humans to develop highly distinct adaptations to the local pressures and environments more quickly than with genetic evolution alone. Robert Boyd and Peter J. Richerson, two strong proponents of cultural evolution, postulate that the act of social learning, or learning in a group as done in group selection, allows human populations to accrue information over many generations.[37] This leads to cultural evolution of behaviors and technology alongside genetic evolution. Boyd and Richerson believe that the ability to collaborate evolved during the Middle Pleistocene, a million years ago, in response to a rapidly changing climate.[37] In 2003, Herbert Gintis examined cultural evolution statistically, offering evidence that societies that promote pro-social norms have higher survival rates than societies that do not.[38][39] Differing evolutionarily stable strategies (ESSs)[edit] The problem with group selection is that for a whole group to get a single trait, it must spread through the whole group first by regular evolution. But, as J. L. Mackie suggested, when there are many different groups, each with a different Evolutionarily Stable Strategy (ESS), there is selection between the different ESSs, since some are worse than others.[40] For example, a group where altruism was universal would outcompete a group where every creature acted in its own interest, but a mixed group of altruists and non-altruists would be vulnerable to cheating by non-altruists within the group.[41] Implications in population biology[edit] Social behaviors such as altruism and group relationships can impact many aspects of population dynamics, such as intraspecific competition and interspecific interactions. In 1871, Darwin argued that group selection occurs when the benefits of cooperation or altruism between subpopulations are greater than the individual benefits of egotism within a subpopulation.[42] This supports the idea of multilevel selection, but kinship also plays an integral role because many subpopulations are composed of closely related individuals. An example of this can be found in lions, which are simultaneously cooperative and territorial.[43] Within a pride, males protect the pride from outside males, and females, who are commonly sisters, communally raise cubs and hunt. However, this cooperation seems to be density dependent. When resources are limited, group selection favors prides that work together to hunt. When prey is abundant, cooperation is no longer beneficial enough to outweigh the disadvantages of altruism, and hunting is no longer cooperative.[43] Interactions between different species can also be affected by multilevel selection. Predator-prey relationships can also be affected. Individuals of certain monkey species howl to warn the group of the approach of a predator.[44] The evolution of this trait benefits the group by providing protection, but could be disadvantageous to the individual if the howling draws the predator's attention to them. By affecting these interspecific interactions, multilevel and kinship selection can change the population dynamics of an ecosystem.[44] Multilevel selection attempts to explain the evolution of altruistic behavior in terms of quantitative genetics. Increased frequency or fixation of altruistic alleles can be accomplished through kin selection, in which individuals engage in altruistic behavior to promote the fitness of genetically similar individuals such as siblings. However, this can lead to inbreeding depression,[45] which typically lowers the overall fitness of a population. However, if altruism were to be selected for through an emphasis on benefit to the group as opposed to relatedness and benefit to kin, both the altruistic trait and genetic diversity could be preserved. However, relatedness should still remain a key consideration in studies of multilevel selection. Experimentally imposed multilevel selection on Japanese quail was more effective by an order of magnitude on closely related kin groups than on randomized groups of individuals.[46]

Criticism[edit] The use of the Price equation to support group selection was challenged by van Veelen in 2012, arguing that it is based on invalid mathematical assumptions.[47] Richard Dawkins and other advocates of the gene-centered view of evolution remain unconvinced about group selection.[48][49][50] In particular, Dawkins suggests that group selection fails to make an appropriate distinction between replicators and vehicles.[51] The psychologist Steven Pinker concluded that "group selection has no useful role to play in psychology or social science", as it "is not a precise implementation of the theory of natural selection, as it is, say, in genetic algorithms or artificial life simulations. Instead it is a loose metaphor, more like the struggle among kinds of tires or telephones."[52] The evolutionary biologist Jerry Coyne summarized the arguments in The New York Times in non-technical terms as follows:[53] Group selection isn't widely accepted by evolutionists for several reasons. First, it's not an efficient way to select for traits, like altruistic behavior, that are supposed to be detrimental to the individual but good for the group. Groups divide to form other groups much less often than organisms reproduce to form other organisms, so group selection for altruism would be unlikely to override the tendency of each group to quickly lose its altruists through natural selection favoring cheaters. Further, little evidence exists that selection on groups has promoted the evolution of any trait. Finally, other, more plausible evolutionary forces, like direct selection on individuals for reciprocal support, could have made humans prosocial. These reasons explain why only a few biologists, like [David Sloan] Wilson and E. O. Wilson (no relation), advocate group selection as the evolutionary source of cooperation.[53]

References[edit] ^ a b Tudge, Colin (31 March 2011). Engineer In The Garden. Random House. p. 115. ISBN 978-1-4464-6698-8.  ^ a b Burkhardt, Richard W. (2005). Patterns of Behavior: Konrad Lorenz, Niko Tinbergen, and the Founding of Ethology. University of Chicago Press. p. 432. ISBN 978-0-226-08090-1.  ^ Wynne-Edwards, V.C. (1962). Animal Dispersion in Relation to Social Behaviour. Edinburgh: Oliver & Boyd. ^ Wynne-Edwards, V. C. (1986) Evolution Through Group Selection, Blackwell. ISBN 0-632-01541-1 ^ Maynard Smith, J. (1964). "Group selection and kin selection". Nature. 201 (4924): 1145–1147. Bibcode:1964Natur.201.1145S. doi:10.1038/2011145a0.  ^ Perrins, Chris. Williams, George C., ed. Survival of Young Swifts in Relation to Brood-Size. Group Selection. Transaction Publishers. pp. 116–118. ISBN 978-0-202-36635-7.  ^ Williams, G.C. (1972) Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought. Princeton University Press.ISBN 0-691-02357-3 ^ Williams, G.C. (editor) (2008) [1971] Group Selection, Transaction Publishers. ISBN 0-202-36222-1 ^ a b Abbot, P. et al. (2011). "Inclusive fitness theory and eusociality". Nature. 471 (7339): E1–E4. doi:10.1038/nature09831. PMID 21430721. CS1 maint: Uses authors parameter (link) CS1 maint: Explicit use of et al. (link) ^ Wade, M. J. (1977). "An experimental study of group selection". Evolution. 31 (1): 134–153. doi:10.2307/2407552. JSTOR 2407552.  ^ Goodnight, C. J.; Stevens, L. (1997). "Experimental studies of group selection: What do they tell us about group selection in nature". American Naturalist. 150: S59–S79. doi:10.1086/286050.  ^ Wade, MJ; Wilson, DS; Goodnight, C; Taylor, D; Bar-Yam, Y; de Aguiar, MA; Stacey, B; Werfel, J; Hoelzer, GA; Brodie ED, 3rd; Fields, P; Breden, F; Linksvayer, TA; Fletcher, JA; Richerson, PJ; Bever, JD; Van Dyken, JD; Zee, P (Feb 18, 2010). "Multilevel and kin selection in a connected world". Nature. 463 (7283): E8–9; discussion E9–10. doi:10.1038/nature08809. PMC 3151728 . PMID 20164866. CS1 maint: Uses authors parameter (link) ^ Koeslag, J.H. (1997). "Sex, the prisoner's dilemma game, and the evolutionary inevitability of cooperation". Journal of Theoretical Biology. 189: 53–61. doi:10.1006/jtbi.1997.0496. PMID 9398503.  ^ Koeslag, J.H. (2003). "Evolution of cooperation: cooperation defeats defection in the cornfield model". Journal of Theoretical Biology. 224: 399–410. doi:10.1016/s0022-5193(03)00188-7.  ^ a b c Wilson, D. S.; Wilson, E. O. (2008). "Evolution 'for the good of the group'". American Scientist. 96 (5): 380–389. doi:10.1511/2008.74.1.  ^ a b c Yang, Jiang-Nan (2013). "Viscous populations evolve altruistic programmed aging in ability conflict in a changing environment". Evolutionary Ecology Research. 15: 527–543.  ^ Wilson, David Sloan (2010). "open letter to Richard Dawkins". Self-published. Retrieved 12 January 2015.  ^ Goodnight, Charles (June 2013). "On multilevel selection and kin selection: Contextual analysis meets direct fitness". Evolution. 67 (6): 1539–1548. doi:10.1111/j.1558-5646.2012.01821.x. PMID 23730749.  ^ Nowak, MA; Tarnita, CE; Wilson, EO (2010). "The evolution of eusociality". Nature. 466: 1057–1062. doi:10.1038/nature09205. PMC 3279739 . PMID 20740005.  ^ Wilson, D.S.; Sober, E. (1994). "Reintroducing group selection to the human behavioral sciences". Behavioral and Brain Sciences. 17 (4): 585–654. doi:10.1017/s0140525x00036104. Archived from the original on 2004-02-03.  ^ a b c O'Gorman, R.; Wilson, D. S.; Sheldon, K. M. (2008). "For the good of the group? Exploring group-level evolutionary adaptations using multilevel selection theory". Group Dynamics: Theory, Research, and Practice. 12 (1): 17–26. doi:10.1037/1089-2699.12.1.17.  ^ Muir, W. M. (2009). "Genetic selection and behaviour". Canadian Journal of Animal Science. 89 (1): 182–182.  ^ Boyd, R.; Richerson, P.J. (2009). "Culture and the evolution of human cooperation". Phil. Trans. R. Soc. B. 364 (1533): 3281–3288. doi:10.1098/rstb.2009.0134.  ^ Wilson, E. O. (2005). "Kin Selection as the Key to Altruism: its Rise and Fall". Social Research. 72 (1): 159–166.  ^ Trivers, R. (1976). "Haploidploidy and the evolution of the social insect". Science. 191 (4224): 250–263. doi:10.1126/science.1108197. PMID 1108197.  ^ a b Wilson, E.O. (1975). Sociobiology: The New Synthesis. Belknap Press. ISBN 0-674-81621-8.  ^ Hamilton, W.D. (1964). "The evolution of social behaviour". Journal of Theoretical Biology. 7 (1): 1–16. doi:10.1016/0022-5193(64)90038-4. PMID 5875341.  ^ West, S.A., Griffin, A.S. & Gardner, A. (2007). "Social semantics: altruism, cooperation, mutualism, strong reciprocity and group selection". Journal of Evolutionary Biology. 20: 415–432. doi:10.1111/j.1420-9101.2006.01258.x. PMID 17305808. CS1 maint: Multiple names: authors list (link) ^ Wilson, David Sloan. "Rethinking the Theoretical Foundation of Sociobiology" (PDF). Retrieved 10 September 2013.  ^ a b Rauch, E. M.; Sayama, H.; Bar-Yam, Y. (2003). "Dynamics and genealogy of strains in spatially extended host-pathogen models". Journal of Theoretical Biology. 221: 655–664. doi:10.1006/jtbi.2003.3127.  ^ Werfel, J.; Bar-Yam, Y. (2004). "The evolution of reproductive restraint through social communication". Proceedings of the National Academy of Sciences of the United States of America. 101 (30): 11019–11020. doi:10.1073/pnas.0305059101. PMC 491990 . PMID 15256603.  ^ Wilson, D.S.; Pollock, G.B.; Dugatkin, L.A (1992). "Can altruism evolve in purely viscous populations?". Evol. Ecol. 6: 331–341. doi:10.1007/bf02270969.  ^ Taylor, P.D. (1992). "Altruism in viscous populations – an inclusive fitness model". Evol. Ecol. 6: 352–356. doi:10.1007/bf02270971.  ^ Mitteldorf, Joshua; Wilson, D.S. (2000). "Population viscosity and the evolution of altruism" (PDF). J. Theor. Biol. 204: 481–496. doi:10.1006/jtbi.2000.2007.  ^ Mesoudi, A.; Danielson, P. (2008). "Ethics, evolution and culture". Theory in Biosciences. 127 (3): 229–240. doi:10.1007/s12064-008-0027-y. PMID 18357481.  ^ Fehr, E.; Fischbacher, Urs (2003). "The nature of human altruism. [Review]". Nature. 425 (6960): 785–791. doi:10.1038/nature02043. PMID 14574401.  ^ a b Boyd, R., & Richerson, P. J. (2009) Culture and the evolution of human cooperation. ^ Gintis, H. (2003). "The hitchhiker's guide to altruism: Gene-culture coevolution, and the internalization of norms". Journal of Theoretical Biology. 220 (4): 407–418. doi:10.1006/jtbi.2003.3104. PMID 12623279.  ^ Gintis, H. (2003). "The hitchhiker's guide to altruism: Gene-culture coevolution, and the internalization of norms". Journal of Theoretical Biology. 220 (4): 407–418. doi:10.1006/jtbi.2003.3104. PMID 12623279.  ^ The selfish Gene (Richard Dawkins) ^ Axelrod, Robert (1984). The Evolution of Cooperation. Basic Books. p. 98. ISBN 978-0465005642.  ^ Darwin,C.: 1871, The Descent of Man, The Heritage Press, New York, 1972. ^ a b Heinsohn, R.; Packer, C. (1995). "Complex cooperative strategies in group-territorialAfrican lions". Science. 269 (5228): 1260–1262. doi:10.1126/science.7652573. PMID 7652573.  ^ a b Cheney, D. L.; Seyfarth, R. M. (1990). How monkeys see the world: Inside the mind of another species. University of Chicago Press. ISBN 978-0-226-10246-7. CS1 maint: Multiple names: authors list (link) ^ Wade, M. J.; Breden, Sept (1981). "Effect of Inbreeding on the Evolution of Altruistic Behavior by Kin Selection". Evolution. 35 (5): 844–858. doi:10.2307/2407855.  ^ Muir, W. M.; et al. (June 2013). "Multilevel selection with kin and non-kin groups,Experimental results with Japanese quail". Evolution. 67 (6): 1598–1606. doi:10.1111/evo.12062. CS1 maint: Explicit use of et al. (link) ^ van Veelen, M; García, J; Sabelis, MW; Egas, M (April 2012). "Group selection and inclusive fitness are not equivalent; the Price equation vs. models and statistics". Journal of Theoretical Biology. 299: 64–80. doi:10.1016/j.jtbi.2011.07.025. PMID 21839750.  ^ See the chapter God's utility function in Dawkins, Richard (1995). River Out of Eden. New York: Basic Books. ISBN 0-465-06990-8.  ^ Dawkins, R. (1994). "Burying the Vehicle Commentary on Wilson & Sober: Group Selection". Behavioural and Brain Sciences. 17 (4): 616–617. Archived from the original on 2006-09-15.  ^ Dennett, D.C. (1994). "E Pluribus Unum? Commentary on Wilson & Sober: Group Selection". Behavioural and Brain Sciences. 17 (4): 617–618. Archived from the original on 2007-12-27.  ^ Richard Dawkins, "Replicators and Vehicles", King's College Sociobiology Group, eds., Current Problems in Sociobiology, Cambridge, Cambridge University Press, (1982), pp. 45–64 ^ Pinker, S. (2012). "The False Allure of Group Selection". Edge. Retrieved June 19, 2012.  ^ a b Coyne, J. A. (2011). Can Darwinism Improve Binghamton?, New York Review of Books, September 9, 2011.

Further reading[edit] Bergstrom, T.C. (2002). "Evolution of Social Behavior: Individual and Group Selection" (PDF). Journal of Economic Perspectives. 16 (2): 67–88. doi:10.1257/0895330027265.  Bijma, P.; Muir, W.M.; Van Arendonk, J.A.M. (2007). "Multilevel Selection 1: Quantitative Genetics of Inheritance and Response to Selection". Genetics. 175 (1): 277–288. doi:10.1534/genetics.106.062711. PMC 1775021 . PMID 17110494.  Bijma, P.; Muir, W.M.; Ellen, E. D.; Wolf, Jason B.; Van Arendonk, J.A.M. (2007). "Multilevel Selection 2: Estimating the Genetic Parameters Determining Inheritance and Response to Selection". Genetics. 175 (1): 289–299. doi:10.1534/genetics.106.062729. PMC 1775010 . PMID 17110493.  Boyd, R.; Richerson, P.J. (2002). "Group Beneficial Norms Spread Rapidly in a Structured Population" (PDF). Journal of Theoretical Biology. 215 (3): 287–296. doi:10.1006/jtbi.2001.2515.  West, S.A.; Griffin, A.S.; Gardner, A. (2008). "Social semantics: how useful has group selection been?". Journal of Evolutionary Biology. 21: 374–385. doi:10.1111/j.1420-9101.2007.01458.x.  Sober, Elliott and Wilson, David Sloan (1998). Unto Others: The Evolution and Psychology of Unselfish Behavior. Cambridge, MA: Harvard University Press. Soltis, J.; Boyd, R.; Richerson, P.J. (1995). "Can Group-functional Behaviors Evolve by Cultural Group Selection? An Empirical Test" (PDF). Current Anthropology. 63: 473–494. doi:10.1086/204381.  Wilson, D. S. (1987). "Altruism in Mendelian populations derived from sibling groups: The haystack model revisited". Evolution. 41 (5): 1059–1070. doi:10.2307/2409191. JSTOR 2409191.  Wilson, David Sloan (2006). Human groups as adaptive units: toward a permanent consensus. In P. Carruthers, S. Laurence & S. Stich (Eds.), The Innate Mind: Culture and Cognition. Oxford: Oxford University Press. Full text (archived from the original on 2009-02-26)

External links[edit] "Altruism and Group Selection". Internet Encyclopedia of Philosophy.  Lloyd, Elisabeth, "Units and Levels of Selection." The Stanford Encyclopedia of Philosophy, (Fall 2005 Edition), Edward N. Zalta (ed.) The Controversy of the Group Selection Theory – a review from the "Science Creative Quarterly" (a blog) Binghamton: D.S. Wilson v t e Sociobiology Topics Behavioural genetics Challenge hypothesis Dual inheritance theory Ethology Evolutionary psychology Evolution of morality Evolutionary models of food sharing Group selection Kin recognition Kin selection Male Warrior hypothesis Reciprocal altruism Sexual selection in human evolution Sex and psychology Sociality (Eusociality, Evolution of eusociality, Presociality) Dunbar's number Supporters Anne Campbell Noam Chomsky Richard Dawkins Daniel Dennett Sarah Blaffer Hrdy Steven Pinker Frans de Waal E. O. Wilson Sociobiology: The New Synthesis Robert Wright Opponents Stephen Jay Gould Leon Kamin Richard Lewontin Steven Rose Not in Our Genes Evolutionary biology portal v t e Evolutionary psychology Processes Altruism Group selection Kin selection Sexual selection Sociobiology Coevolution Evolutionarily stable strategy Areas Psychological development Morality Religion Depression Developmental psychopathology Educational psychology Evolutionary aesthetics Music Darwinian literary studies Evolution of emotion Biologists/ Neuroscientists John Crook Charles Darwin Richard Dawkins Jared Diamond W. D. Hamilton Peter Kropotkin Gordon Orians Jaak Panksepp Margie Profet Giacomo Rizzolatti Randy Thornhill Robert Trivers Carel van Schaik Claus Wedekind Wolfgang Wickler David Sloan Wilson E. O. Wilson George C. Williams Anthropologists Jerome H. Barkow Robert Boyd Robin Dunbar Daniel Fessler Mark Flinn Joseph Henrich Ruth Mace Daniel Nettle Frank Salter Stephen Shennan Donald Symons John Tooby Behavioral economists/ Political scientists Samuel Bowles Ernst Fehr Herbert Gintis Dominic D. P. Johnson Gad Saad Literary theory/ Aesthetics Joseph Carroll Denis Dutton Psychologists/ Cognitive scientists Simon Baron-Cohen Justin L. Barrett Jay Belsky David F. Bjorklund Paul Bloom Pascal Boyer Joseph Bulbulia David Buss Josep Call Anne Campbell Leda Cosmides Martin Daly Daniel Dennett Paul Ekman Anne Fernald David C. Geary Gerd Gigerenzer Jonathan Haidt Judith Rich Harris Aurelio José Figueredo Stephen Kaplan Douglas T. Kenrick Simon M. Kirby Robert Kurzban Michael T. McGuire Geoffrey Miller Randolph M. Nesse Steven Neuberg David Perrett Steven Pinker Paul Rozin Mark Schaller David P. Schmitt Todd K. Shackelford Roger Shepard Peter K. Smith Dan Sperber Anthony Stevens Frank Sulloway Michael Tomasello Mark van Vugt Glenn Wilson Margo Wilson Related subjects and articles The Adapted Mind The Evolution of Human Sexuality Evolutionary Psychology Evolution and Human Behavior Memetics Sociobiology Evolutionary neuroscience Human evolution Sociocultural evolution Evolutionary anthropology Evolutionary medicine Evolutionary linguistics Evolutionary psychology and culture Biosocial criminology Criticism of evolutionary psychology Lists Evolutionary psychologists Evolutionary psychology research groups and centers Bibliography of evolution and human behavior Evolutionary psychology Psychology portal Evolutionary biology portal v t e Eusociality Topics Evolution of eusociality Presociality Social insects Gamergate Group selection Haplodiploidy Identity in social insects Kin recognition Kin selection Sexual selection in social insects Thelytoky Worker policing Groups Hymenoptera Ant Apidae Crabronidae Halictidae Honey bee Vespidae Mammalia Blesmol Dwarf mongoose Meerkat Crustacea Synalpheus Thysanoptera Kladothrips Hemiptera Aphididae Coleoptera Austroplatypus incompertus Isoptera In culture Bee (mythology) Pioneers, works Karl von Frisch The Dancing Bees 1927 Charles Duncan Michener The Bees of the World 2000 E. O. Wilson The Ants 1990 Sociobiology: The New Synthesis 1975 Retrieved from "" Categories: SelectionPopulation geneticsHidden categories: CS1 maint: Uses authors parameterCS1 maint: Explicit use of et al.CS1 maint: Multiple names: authors list

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