Group selection

Group selection is a proposed mechanism of evolution in which

natural selection acts at the level of the group, instead of at the
more conventional level of the individual.

Early authors such as V. C. Wynne-Edwards and Konrad Lorenz

argued that the behavior of animals could affect their survival and
reproduction as groups, speaking for instance of actions for the
good of the species. In the 1930s, R.A. Fisher and J.B.S. Haldane
proposed the concept of kin selection, arguing that animals should
sacrifice for their relatives, and thereby implying that they should
not sacrifice for non-relatives; Haldane even jokingly introduced a
Early explanations of social behavior, such as the
mathematical basis for familial altruism, suggesting that he would
lekking of blackcock, spoke of "the good of the
die for two siblings or eight cousins. From the mid 1960s, species".[1] Blackcocks at the Lek watercolour
evolutionary biologists such as John Maynard Smith argued that and bodycolour by Archibald Thorburn, 1901.
natural selection acted primarily at the level of the individual. They
argued on the basis of mathematical models that individuals would
not altruistically sacrifice fitness for the sake of a group. They persuaded the majority of biologists that group selection did not
occur, other than in special situations such as the haplodiploid social insects like honeybees (in the Hymenoptera), where kin
selection was possible.

In 1994 David Sloan Wilson and Elliott Sober argued for multi-level selection, including group selection, on the grounds that
groups, like individuals, could compete. In 2010 three authors including E. O. Wilson, known for his work on social insects
especially ants, again revisited the arguments for group selection. They argued that group selection can occur when competition
between two or more groups, some containing altruistic individuals who act cooperatively together, is more important for survival
than competition between individuals within each group. Howard Rachlin, William M. Baum, and Carsta Simon have proposed
group selection of behavioral patterns during ontogeny parallel to group selection during phylogeny.[2][3][4] Their proposals
provoked a strong rebuttal from a large group of evolutionary biologists and behavior analysts.[5]

Contents
Early developments
Kin selection and inclusive fitness theory
Multilevel selection theory
Applications
Differing evolutionarily stable strategies
Implications in population biology
Gene-culture coevolution in humans
Criticism
References
Further reading
External links
Early developments
Charles Darwin developed the theory of evolution in his book, Origin of Species. Darwin also made the first suggestion of group
selection in The Descent of Man that the evolution of groups could affect the survival of individuals. He wrote, "If one man in a
tribe... invented a new snare or weapon, the tribe would increase in number, spread, and supplant other tribes. In a tribe thus
rendered more numerous there would always be a rather better chance of the birth of other superior and inventive members."[6][7]

Once Darwinism had been accepted in the modern synthesis of the mid-twentieth century, 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][8] without
actually studying survival value in the field.[8] Richard Dawkins noted that Lorenz was a "'good of the species' man"[9] so
accustomed to group selection thinking that he did not realize his views "contravened orthodox Darwinian theory".[9] 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.[8] In 1962, group selection was used as a popular explanation for adaptation by the zoologist V. C.
Wynne-Edwards.[10][11] In 1976, Richard Dawkins wrote a well-known book on the importance of evolution at the level of the
gene or the individual, The Selfish Gene.[12]

From the mid 1960s, evolutionary biologists argued that natural selection acted primarily
at the level of the individual. In 1964, John Maynard Smith,[13] C. M. Perrins (1964),[14]
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.[15][16]

It was at that time generally agreed that the primary exception of social group selection
was in the social insects, and the explanation was limited to the unique inheritance system
(involving haplodiploidy) of the eusocial Hymenoptera such as honeybees, which
encourages kin selection, since workers are closely related.[5]

Kin selection and inclusive fitness theory

Social behavior in
Early group selection models assumed that genes acted independently, for example a gene honeybees is explained by
that coded for cooperation or altruism. Genetically-based reproduction of individuals kin selection: their
implies that, in group formation, the altruistic genes would need a way to act for the haplodiploid inheritance
benefit of members in the group to enhance the fitness of many individuals with the same system makes workers very
gene.[17] But it is expected from this model that individuals of the same species would closely related to their
queen (centre).
compete against each other for the same resources. This would put cooperating individuals
at a disadvantage, making genes for cooperation likely to be eliminated. Group selection
on the level of the species is flawed because it is difficult to see how selective pressures would be applied to competing/non-
cooperating individuals.[12]

Experiments from the late 1970s suggested that selection involving groups was possible.[18] Kin selection between related
individuals is accepted as an explanation of altruistic behavior. In this model, genetically related individuals cooperate because
survival advantages to one individual also benefit kin who share some fraction of the same genes, giving a mechanism for
favoring genetic selection.[19]

Inclusive fitness theory, first proposed by W. D. Hamilton in the early 1960s, gives a selection criterion for evolution of social
traits when social behavior is costly to an individual organism's survival and reproduction. This behavior could emerge under
conditions such that the statistical likelihood that benefits accrue to the survival and reproduction of other organisms whom also
carry the social trait. Inclusive fitness theory is a general treatment of the statistical probabilities of social traits accruing to any
other organisms likely to propagate a copy of the same social trait. Kin selection theory treats the narrower but simpler case of the
benefits to close genetic relatives (or what biologists call 'kin') who may also carry and propagate the trait. A significant group of
biologists support inclusive fitness as the explanation for social behavior in a wide range of species, as supported by experimental
data. An article was published in Nature with over a hundred coauthors.[5]

One of the questions about kin selection is the requirement that individuals must know if other individuals are related to them, or
kin recognition. Any altruistic act has to preserve similar genes. One argument given by Hamilton is that many individuals
operate in "viscous" conditions, so that they live in physical proximity to relatives. Under these conditions, they can act
altruistically to any other individual, and it is likely that the other individual will be related. 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.[20] and Taylor[21] 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. Mitteldorf and D. S. Wilson 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.[22] By assuming individual differences in adaptations, Yang 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.[23]

Another explanation for the recognition of genes for altruism is that a single trait, group reciprocal kindness, is capable of
explaining the vast majority of altruism that is generally accepted as "good" by modern societies. The phenotype of altruism
relies on recognition of the altruistic behavior by itself. The trait of kindness will be recognized by sufficiently intelligent and
undeceived organisms in other individuals with the same trait. Moreover, the existence of such a trait predicts a tendency for
kindness to unrelated organisms that are apparently kind, even if the organisms are of a completely different species. The gene
need not be exactly the same, so long as the effect or phenotype is similar. Multiple versions of the gene—or even meme—would
have virtually the same effect. This explanation was given by Richard Dawkins as an analogy of a man with a green beard.
Green-bearded men are imagined as tending to cooperate with each other simply by seeing a green beard, where the green beard
trait is incidentally linked to the reciprocal kindness trait.[12]

Multilevel selection theory

Kin selection or inclusive fitness is accepted as an explanation for cooperative behavior in many species, but there are some
species, including some human behavior, that are difficult to explain with only this approach. In particular, it does not seem to
explain the rapid rise of human civilization. David Sloan Wilson has argued that other factors must also be considered in
evolution.[24] Wilson and others have continued to develop group selection models.[25]

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.[17] 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.[19][26] 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".[27]

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.[28]
In 2010, M. A. Nowak, C. E. Tarnita and 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.[29] A response from 137 other evolutionary biologists argued
"that their arguments are based upon a misunderstanding of evolutionary theory and a misrepresentation of the empirical
literature".[5]

Wilson compared the layers of competition and evolution to nested

sets of Russian matryoshka dolls.[30] 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-benefiting trait to spread.[31]

Multilevel selection theory focuses on the phenotype because it

looks at the levels that selection directly acts upon.[30] For humans,
social norms can be argued to reduce individual level variation and
competition, thus shifting selection to the group level. The David Sloan Wilson and Elliott Sober's 1994
assumption is that variation between different groups is larger than Multilevel Selection Model, illustrated by a nested
variation within groups. Competition and selection can operate at set of Russian matryoshka dolls. Wilson himself
compared his model to such a set.
all levels regardless of scale. Wilson wrote, "At all scales, there
must be mechanisms that coordinate the right kinds of action and
prevent disruptive forms of self-serving behavior at lower levels of social organization."[24] E. O. Wilson summarized, "In a
group, selfish individuals beat altruistic individuals. But, groups of altruistic individuals beat groups of selfish individuals."[32]

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.[31]

MLS theory can be used to evaluate the balance between group selection and individual selection in specific cases.[31] 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.[33] 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.[34]

Wilson and Sober's work revived interest in multilevel selection. In a 2005 article,[35] 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.[36] 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 Sociobiology: A New Synthesis was published in
1975.[37] These 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:[38]

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,[39] 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.[40]

Spatial populations of predators and prey show restraint of reproduction at equilibrium, both individually and through social
communication, 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.[41][42]

Rauch et al.'s analysis[41] of host-parasite evolution, which even E. O. Wilson recognised as a situation where group selection
was possible (1975), is broadly hostile to group selection. 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."[37]

Applications

Differing evolutionarily stable strategies

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, there is selection between the different strategies, since some are worse than others.[43] For example, a group
where altruism was universal would indeed outcompete a group where every creature acted in its own interest, so group selection
might seem feasible; but a mixed group of altruists and non-altruists would be vulnerable to cheating by non-altruists within the
group, so group selection would collapse.[44]

Implications in population biology

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.[6] 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.[45] 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.[45]

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.[46] 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.[46]
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,[47] 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.[48]

Gene-culture coevolution in humans

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.[49] It treats culture as a separate evolutionary
system that acts in parallel to the usual genetic evolution to transform human traits.[50] 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.[51]

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 Humanity has developed
populations to accrue information over many generations.[52] This leads to cultural extremely rapidly, arguably
evolution of behaviors and technology alongside genetic evolution. Boyd and Richerson through gene-culture
believe that the ability to collaborate evolved during the Middle Pleistocene, a million coevolution, leading to
complex cultural artefacts
years ago, in response to a rapidly changing climate.[52]
like the gopuram of the Sri
Mariammam temple,
In 2003, the behavioral scientist Herbert Gintis examined cultural evolution statistically,
Singapore.
offering evidence that societies that promote pro-social norms have higher survival rates
than societies that do not.[53] Gintis wrote that genetic and cultural evolution can work
together. Genes transfer information in DNA, and cultures transfer information encoded in brains, artifacts, or documents.
Language, tools, lethal weapons, fire, cooking, etc., have a long-term effect on genetics. For example, cooking led to a reduction
of size of the human gut, since less digestion is needed for cooked food. Language led to a change in the human larynx and an
increase in brain size. Projectile weapons led to changes in human hands and shoulders, such that humans are much better at
throwing objects than the closest human relative, the chimpanzee.[54]

Criticism
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.[55]

Richard Dawkins and other advocates of the gene-centered view of evolution remain unconvinced about group
selection.[56][57][58] In particular, Dawkins suggests that group selection fails to make an appropriate distinction between
replicators and vehicles.[59]
The psychologist Steven Pinker concluded that "group selection has no useful role to play in psychology or social science", since
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."[60]

The evolutionary biologist Jerry Coyne summarized the arguments in The New York Review of Books in non-technical terms as
follows:[61]

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.[61]

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Further reading
Bergstrom, T. C. (2002). "Evolution of Social Behavior: Individual and Group Selection" (http://www.econ.ucsb.ed
u/~tedb/Evolution/groupselection.pdf) (PDF). Journal of Economic Perspectives. 16 (2): 67–88.
CiteSeerX 10.1.1.377.5059 (https://citeseerx.ist.psu.edu/viewdoc/summary?doi=10.1.1.377.5059).
doi:10.1257/0895330027265 (https://doi.org/10.1257%2F0895330027265).
Bijma, P.; Muir, W. M.; Van Arendonk, J. A. M. (2007). "Multilevel Selection 1: Quantitative Genetics of
Inheritance and Response to Selection" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1775021). Genetics. 175
(1): 277–288. doi:10.1534/genetics.106.062711 (https://doi.org/10.1534%2Fgenetics.106.062711).
PMC 1775021 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1775021). PMID 17110494 (https://www.ncbi.nlm.
nih.gov/pubmed/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" (https://www.ncbi.nlm.ni
h.gov/pmc/articles/PMC1775010). Genetics. 175 (1): 289–299. doi:10.1534/genetics.106.062729 (https://doi.org/
10.1534%2Fgenetics.106.062729). PMC 1775010 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1775010).
PMID 17110493 (https://www.ncbi.nlm.nih.gov/pubmed/17110493).
Boyd, R.; Richerson, P. J. (2002). "Group Beneficial Norms Spread Rapidly in a Structured Population" (http://ww
w.sscnet.ucla.edu/anthro/faculty/boyd/BoydRichersonJTB2002_Clean.pdf) (PDF). Journal of Theoretical Biology.
215 (3): 287–296. CiteSeerX 10.1.1.405.1548 (https://citeseerx.ist.psu.edu/viewdoc/summary?doi=10.1.1.405.15
48). doi:10.1006/jtbi.2001.2515 (https://doi.org/10.1006%2Fjtbi.2001.2515). PMID 12054837 (https://www.ncbi.nl
m.nih.gov/pubmed/12054837).
West, S. A.; Griffin, A. S.; Gardner, A. (2008). "Social semantics: how useful has group selection been?" (http://w
ww3.interscience.wiley.com/cgi-bin/fulltext/119412422/PDFSTART). Journal of Evolutionary Biology. 21: 374–
385. doi:10.1111/j.1420-9101.2007.01458.x (https://doi.org/10.1111%2Fj.1420-9101.2007.01458.x).
Sober, Elliott; Wilson, David Sloan (1998). Unto Others: The Evolution and Psychology of Unselfish Behavior.
Harvard University Press. ISBN 978-0674930476.
Soltis, J.; Boyd, R.; Richerson, P. J. (1995). "Can Group-functional Behaviors Evolve by Cultural Group
Selection? An Empirical Test" (http://www.des.ucdavis.edu/faculty/richerson/SoltisBoydRichersonCA95.pdf)
(PDF). Current Anthropology. 63 (3): 473–494. doi:10.1086/204381 (https://doi.org/10.1086%2F204381).
Wilson, David Sloan (1987). "Altruism in Mendelian populations derived from sibling groups: The haystack model
revisited". Evolution. 41 (5): 1059–1070. doi:10.2307/2409191 (https://doi.org/10.2307%2F2409191).
JSTOR 2409191 (https://www.jstor.org/stable/2409191).
Wilson, David Sloan (2006). P. Carruthers, S. Laurence & S. Stich (ed.). Human groups as adaptive units: toward
a permanent consensus (https://web.archive.org/web/20090226134728/http://evolution.binghamton.edu/dswilso
n/resources/publications_resources/DSW02.pdf) (PDF). The Innate Mind: Culture and Cognition. Oxford
University Press. ISBN 978-0195310146. Archived from the original (http://evolution.binghamton.edu/dswilson/re
sources/publications_resources/DSW02.pdf) (PDF) on 26 February 2009.

External links
"Altruism and Group Selection" (http://www.iep.utm.edu/altr-grp). Internet Encyclopedia of Philosophy.
Lloyd, Elisabeth, "Units and Levels of Selection." (http://plato.stanford.edu/archives/fall2005/entries/selection-unit
s/) The Stanford Encyclopedia of Philosophy, (Fall 2005 Edition), Edward N. Zalta (ed.)
The Controversy of the Group Selection Theory (http://www.scq.ubc.ca/?p=170) – a review from the "Science
Creative Quarterly" (a blog)
Binghamton: D.S. Wilson (https://web.archive.org/web/20070715062538/http://evolution.binghamton.edu/dswilso
n/publications.html)

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