Group selection

In evolutionary biology, group selection refers to the idea that alleles can become fixed or spread in a population because of the benefits they bestow on groups, regardless of the alleles' effect on the fitness of individuals within that group.

Group selection was used as a popular explanation for adaptations, especially by V. C. Wynne-Edwards^[1][2]. For several decades, however, critiques, particularly by George C. Williams^[3][4], John Maynard Smith^[5] and C.M. Perrins (1964), cast serious doubt on group selection as a major mechanism of evolution, and only recently have group selection models seen a resurgence (albeit not as a fundamental mechanism but as a phenomenon emergent from standard selection).

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Overview

Specific syndromes of selective factors can create situations in which groups are selected because they display group properties which are selected-for. Some mosquito-transmitted rabbit viruses, for instance, are only transmitted to uninfected rabbits from infected rabbits which are still alive. This creates a selective pressure on every group of viruses already infecting a rabbit not to become too virulent and kill their host rabbit before enough mosquitoes have bitten it. In natural systems such viruses display much lower virulence levels than do mutants of the same viruses that in laboratory culture readily out-compete non-virulent variants (or than do tick-transmitted viruses—ticks, unlike mosquitoes, bite dead rabbits).

However, theoretical models of the 1960s seemed to imply that the effect of group selection was negligible. Alleles are likely to be held on a population-wide level, leaving nothing for group selection to select for. Additionally, generation time is much longer for groups than it is for individuals. Assuming conflicting selection pressures, individual selection will occur much faster, swamping any changes potentially favored by group selection. The Price equation can partition variance caused by natural selection at the individual level and the group level, and individual level selection generally causes greater effects.

Experimental results starting in the late 1970s demonstrated that group selection was far more effective than theoretical models ever would have predicted^[6]. A review of this experimental work has shown that the early group selection models were flawed because they assumed that genes acted independently, whereas in the experimental work it was apparent that gene interaction, and more importantly, genetically based interactions among individuals, were an important source of the response to group selection (e.g. ^[7]). As a result many are beginning to recognize that group selection, or more appropriately multilevel selection, is potentially an important force in evolution.

More recently, Yaneer Bar-Yam has claimed that the gene-centered view (and thus Fisher's treatment of evolution) relies upon a mathematical approximation that is not generally valid. Bar-Yam argues that the approximation is a dynamic form of the Mean Field approximation frequently used in physics and whose limitations are recognized there. In biology, the approximation breaks down when there are spatial populations resulting in inhomogeneous genetic types (called symmetry breaking in physics). Such symmetry breaking may also correspond to speciation.

Spatial populations of predators and prey have also been shown to show restraint of reproduction at equilibrium, both individually^[8] and through social communication^[9], 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^[8], 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.^[10]

Multilevel selection theory

See also: Unit of selection

In recent years, the limitations of earlier models have been addressed, and newer models suggest that selection may sometimes act above the gene level. Recently David Sloan Wilson and Elliot Sober have argued that the case against group selection has been overstated. They focus their argument on whether groups can have functional organization in the same way individuals do and, consequently, if groups can also be "vehicles" for selection. For example, groups that cooperate better may have out-reproduced those which did not. Resurrected in this way, Wilson & Sober's new group selection is usually called multilevel selection theory.^[11]

Although Richard Dawkins and fellow advocates of the gene-centered view of evolution remain unconvinced (see, for example, ^[12][13][14]), Wilson & Sober's work has been part of a broad revival of interest in multilevel selection as an explanation for evolutionary phenomena. Indeed, in a 2005 article^[15], E. O. Wilson (often regarded as the father of sociobiology) argued that kin selection could no longer be thought of as underlying the evolution of extreme sociality, for two reasons. First, some authors have shown that the argument that haplodiploid inheritance, characteristic of the Hymenoptera, creates a strong selection pressure towards nonreproductive castes is mathematically flawed (e.g. ^[16]). Secondly, 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^[10], including a variety of insect species, as well as a rodent species (the naked mole rat). Wilson suggests the equation for Hamilton's rule:^[17]

(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

in which b_k is the benefit to kin (b in the original equation) and b_e 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 b_e>>rb_k, so that altruism needs to be explained in terms of selection at the colony level rather than at the kin level.

However, even more recently, Foster et al. (2006) have argued that Wilson made a series of basic errors in logic in making these arguments and that kin selection and group selection are, in fact, not in opposition at all.^[18]

Reeve & Hölldobler (2007) have further expanded upon the group selection model, with a new "superorganism" model, in which groups composed of individuals that invest more in between-group competition will be favored over groups composed of individuals that invest more in within-group competition.^[19]

References

1. ^ Wynne-Edwards, V.C. (1962). Animal Dispersion in Relation to Social Behaviour. Edinburgh: Oliver & Boyd.
2. ^ Wynne-Edwards, V. C. (1986) Evolution Through Group Selection, Blackwell. ISBN 0-632-01541-1
3. ^ Williams, G.C. (1972) Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought. Princetown UP.ISBN 0-691-02357-3
4. ^ Williams, G.C. (1986) Evolution Through Group Selection. Blackwell. ISBN 0-632-01541-1
5. ^ Maynard Smith, J. (1964) Group selection and kin selection Nature 201:1145–1147
6. ^ Wade, M. J. 1977. An experimental study of group selection. Evolution 31:134–153
7. ^ Goodnight, C. J. and L. Stevens. 1997. Experimental studies of group selection: What do they tell us about group selection in nature. American Naturalist 150:S59–S79.
8. ^ ^{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.
9. ^ 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, 11019–11020.
10. ^ ^{a b} Wilson, E.O. 1975. Sociobiology: The New Synthesis Belknap Press, ISBN 0-674-81621-8.
11. ^ link Wilson, D.S. & Sober, E. 1994. Reintroducing group selection to the human behavioral sciences. Behavioral and Brain Sciences 17 (4): 585–654.
12. ^ See the chapter God's utility function in Dawkins, Richard (1995). River Out of Eden. New York: Basic Books. ISBN 0-465-06990-8. 
13. ^ link Dawkins, R. (1994). Burying the Vehicle. Commentary on Wilson & Sober: Group Selection. Behavioural and Brain Sciences. 17 (4): 616–617.
14. ^ link Dennett, D.C. (1994). E Pluribus Unum? Commentary on Wilson & Sober: Group Selection. Behavioural and Brain Sciences. 17 (4): 617–618.
15. ^ Wilson, E. O. (2005). Kin Selection as the Key to Altruism: its Rise and Fall. "Social Research" 72 (1): 159–166.
16. ^ Trivers, R. (1979) Science 191(4224), 250-263
17. ^ Hamilton, W.D. (1964) The evolution of social behavior Journal of Theoretical Biology 1:295–311.
18. ^ Foster, KR, Wenseleers T, and Ratnieks FLW 2006. Kin selection is the key to altruism. Trends in Ecology and Evolution, 21:57–60
19. ^ Reeve, H.K. and Hölldobler, B. 2007. The emergence of a superorganism through intergroup competition. Proceedings of the National Academy of Sciences 104: 9736-9740

Further reading

• Bergstrom, T.C. (2002). Evolution of Social Behavior: Individual and Group Selection, Journal of Economic Perspectives, 16, 67-88. Full text

• Bijma, P., Muir, W.M. & Van Arendonk, J.A.M. (2007). Multilevel Selection 1: Quantitative Genetics of Inheritance and Response to Selection. Genetics 175: 277-288 link

• Bijma, P., Muir, W.M., Ellen, E. D., Jason B. Wolf, J. B. & Van Arendonk, J.A.M. (2007). Multilevel Selection 2: Estimating the Genetic Parameters Determining Inheritance and Response to Selection. Genetics 175: 289-299 link

• Boyd, R. & Richerson, P.J. (2002). Group Beneficial Norms Spread Rapidly in a Structured Population. Journal of Theoretical Biology, 215, 287–296. Full text

• Soltis, J., Boyd, R. & Richerson, P.J.(1995). Can Group-functional Behaviors Evolve by Cultural Group Selection? An Empirical Test. Current Anthropology, 63, 473–494. Full text

• Wilson, D.S. (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

See also

• Eva Jablonka

• Link to publications by D.S. Wilson