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Multilevel Selection Theory and Major Evolutionary Transitions

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Abstract

The concept of a group as comparable to a single organism has had a long and turbulent history. Currently, methodological individualism dominates in many areas of psychology and evolution, but natural se-lection is now known to operate at multiple levels of the biological hierarchy. When between-group selection dom-inates within-group selection, a major evolutionary tran-sition occurs and the group becomes a new, higher-level organism. It is likely that human evolution represents a major transition, and this has wide-ranging implications for the psychological study of group behavior, cognition, and culture. The concept of a social group as a single organism has a long history in scientific and intellectual thought. According to Daniel Wegner (1986, p. 185), Social commentators once found it very useful to analyze the be-havior of groups by the same expedient used in analyzing the be-havior of individuals. The group, like the person, was assumed to be sentient, to have a form of mental activity that guides action. Rousseau and Hegel were the early architects of this form of analysis, and it became so widely used in the 19th and early 20th centuries that almost every early social theorist we now recognize as a contributor to modern social psychology held a similar view. Nevertheless, during the second half of the 20th century this view of society was eclipsed by a more reductionistic and indi-vidualistic view. Donald Campbell (1994, p. 23) wrote: ''Meth-odological individualism dominates our neighboring fields of
Multilevel Selection Theory and
Major Evolutionary Transitions
Implications for Psychological Science
David Sloan Wilson,
1
Mark Van Vugt,
2
and Rick O’Gorman
3
1
Departments of Biology and Anthropology, Binghamton University;
2
University of Kent at Canterbury, Canterbury,
United Kingdom; and
3
Sheffield Hallam University, Sheffield, United Kingdom
ABSTRACT—The concept of a group as comparable to a
single organism has had a long and turbulent history.
Currently, methodological individualism dominates in
many areas of psychology and evolution, but natural se-
lection is now known to operate at multiple levels of the
biological hierarchy. When between-group selection dom-
inates within-group selection, a major evolutionary tran-
sition occurs and the group becomes a new, higher-level
organism. It is likely that human evolution represents a
major transition, and this has wide-ranging implications
for the psychological study of group behavior, cognition,
and culture.
KEYWORDS—group selection; human evolution; multilevel
selection theory; group psychology; culture
The concept of a social group as a single organism has a long
history in scientific and intellectual thought. According to
Daniel Wegner (1986, p. 185),
Social commentators once found it very useful to analyze the be-
havior of groups by the same expedient used in analyzing the be-
havior of individuals. The group, like the person, was assumed to
be sentient, to have a form of mental activity that guides action.
Rousseau and Hegel were the early architects of this form of
analysis, and it became so widely used in the 19th and early 20th
centuries that almost every early social theorist we now recognize
as a contributor to modern social psychology held a similar view.
Nevertheless, during the second half of the 20th century this
view of society was eclipsed by a more reductionistic and indi-
vidualistic view. Donald Campbell (1994, p. 23) wrote: ‘‘Meth-
odological individualism dominates our neighboring fields of
economics, much of sociology, and all of psychology’s excursions
into organizational theory. This is the dogma that all human
social group processes are to be explained by laws of individual
behavior.’’
Developments in evolutionary biology seemed to affirm the
individualistic turn in psychology. Darwin wrote about how
groups can potentially, but not invariably, evolve into adaptive
units (Richards, 1987). Unfortunately, many of his followers
assumed that natural selection operates on individuals, groups,
species, and ecosystems, as though there were no need to dis-
tinguish among levels of the biological hierarchy. These ideas
were criticized in the 1960s, and a two-part consensus emerged
(Williams 1966). First, higher-level entities such as social
groups can evolve into adaptive units, but only by a process of
higher-level selection. For example, an altruistic behavior that
benefits others at the expense of the self is selectively disad-
vantageous within groups. However, if there are many groups in
the total population that vary in the frequency of altruists, the
most altruistic groups will differentially contribute to the total
gene pool. Between-group selection favors altruism and can
counteract within-group selection if it is sufficiently strong,
causing the altruistic trait to evolve in the total population. This
way of conceptualizing evolution is called multilevel selection
(MLS) theory (Sober & Wilson, 1998).
Even though group-level adaptations can evolve in theory, the
second part of the consensus concluded that they seldom do so
in the real world, because group-level selection is almost
invariably weaker than individual-level selection. This con-
clusion was so widely accepted that group selection became a
pariah concept, taught primarily as an example of how not to
think. The theoretical justification for individualism in psy-
chology seemed secure.
Nevertheless, much has happened in evolutionary biology
during the last half century (Wilson & Wilson, 2007). The first
part of the 1960s consensus remains valid: Adaptations at any
given level of the biological hierarchy require a process of
Address correspondence to David Sloan Wilson, Department of Bi-
ology, Binghamton University, Binghamton, New York, 13902-6000;
e-mail: dwilson@binghamton.edu.
CURRENT DIRECTIONS IN PSYCHOLOGICAL SCIENCE
6Volume 17—Number 1Copyright r2008 Association for Psychological Science
natural selection at that level and tend to be undermined by
lower levels of selection. The second part of the consensus has
proven to be erroneous: Higher-level selection can be a sig-
nificant evolutionary force, one that sometimes even dominates
lower-level selection, causing the higher-level unit to become an
organism in every sense of the word. Ironically, given group
selection’s previous pariah status, it is now the concept of groups
as organisms that stands on a firm scientific foundation. More-
over, it is likely that human evolution represents such an evo-
lutionary transition, and this has profound implications for
psychology and all other human-related subjects.
ORGANISMS AS GROUPS
When between-group selection dominates within-group selec-
tion, a major evolutionary transition occurs. The social group
becomes a higher-level organism and the members of the group
acquire an organ-like status. This idea was first proposed to
explain the evolution of eukaryotic (nucleated) cells, not by
small mutational steps from prokaryotic (bacterial) cells but as
highly integrated symbiotic associations of bacteria. The idea
was then generalized to include other major transitions, in-
cluding the first cells, multicellular organisms, social insect
colonies, and even the origin of life as groups of cooperating
molecular interactions (Maynard Smith & Szathmary, 1995).
Major transitions have a number of hallmarks: First, they are
rare events in the history of life. It is not easy for between-group
selection to dominate within-group selection. All species of
eusocial insects (e.g., ants, bees, wasps, and termites), for ex-
ample, are thought to be derived from only 15 original transitions
from solitary insect species. Second, major transitions have
momentous consequences once they occur. Individuals and
uncoordinated groups are no match for the new superorganisms,
which quickly become ecologically dominant. Third, the tran-
sition is never complete. Even multicellular organisms, which
might seem like paradigms of internal harmony, contain a dis-
turbing number of genes that spread at the expense of other
genes in the same organism rather than for the good of the
organism (cf. intragenomic conflict).
THE HUMAN MAJOR TRANSITION: IMPLICATIONS FOR
PSYCHOLOGICAL SCIENCE
It is likely that early human evolution represented a major
transition, turning our ancestral groups into the primate equiv-
alent of bodies or beehives. All of the hallmarks are present:
It was a rare event, occurring only once among primates. The
consequences were momentous; mere individuals and less co-
ordinated groups were no match for the new superorganisms,
which spread over the globe, eliminating other hominid species
and thousands of other species along the way. The transition is
not complete; individuals still succeed at the expense of other
individuals within the same group. The scope for within-group
selection is merely suppressed, turning between-group selection
into a relatively stronger evolutionary force.
This multilevel view of human evolution, with a strong (but not
exclusive) emphasis on group selection, has foundational im-
plications for psychological science. These implications are not
entirely new, however, because psychology has its own tradition
of group-level thinking, as we stressed at the beginning of this
article. Instead, MLS theory can provide a new foundation
for longstanding themes in psychological research, a sample of
which will now be described.
Psychology of Altruism, Cooperation, and Morality
A major transition requires mechanisms that suppress conflict
among individuals within groups, enabling groups to become the
primary unit of selection. Multicellular organisms and social
insect colonies could not function as adaptive units without in-
ternal social-control mechanisms (Maynard Smith & Szathmary,
1995). In humans, the traits associated with moral systems ap-
pear to perform the same function. Small-scale human society
(the only scale during most of human evolution) is remarkable for
the degree of social control that members can exert over each
other. In human societies, unlike those of most primate species,
no single individual can dominate the others in their group.
Behaviors that benefit some members at the expense of others are
easily detected, communicated, and punished at relatively low
cost to the punishers. These social-control mechanisms estab-
lish a kind of guarded egalitarianism that Boehm (1999) terms a
‘‘moral community,’’ and which characterizes virtually all known
hunter-gatherer societies.
The experimental games literature shows the importance of
social control for the maintenance of cooperation in human
groups. When given an opportunity to cooperate, most people are
moderately generous until they perceive that they are being
exploited by more selfish ingroup members, after which they
withhold their own cooperation (De Cremer & Van Vugt, 1999).
At least some members are highly motivated to punish selfish
behavior, however, even at their own private expense, resulting
in high levels of cooperation (Fehr & Ga
¨chter, 2002). Group-
level selection thinking forces researchers to reconsider the
notion of Homo economicus and replace it with a more complex
picture, one that includes human preferences for altruism,
benevolence, retaliation, contrition, fairness, forgiveness, and so
on.
These and other traits associated with human morality and
cooperation are based on neurobiological mechanisms that are
primarily automatic and emotive (e.g., social emotions like anger
and guilt) rather than conscious and deliberative. Moral intu-
ition comes first and is only partially overridden by moral rea-
soning. Haidt (2007) shows that early theorizing about morality,
dating back to first psychologists like Wilhelm Wundt and
William James, can be placed on a contemporary foundation
based on MLS theory.
Volume 17—Number 1 7
David Sloan Wilson, Mark Van Vugt, and Rick O’Gorman
Group Cognition and Performance
Cooperation evolves in the context of cognitive activities such as
perception, attention, memory, and decision making, in addition
to physical activities such as hunting, gathering, warfare, and
childcare. The social insects are well known to cooperate on
cognitive tasks, to the point where they can truly be said to
possess a group mind (Seeley, 1995). Just as individual cognition
is based on interactions between neurons, with any particular
neuron playing a limited role, group cognition is based on social
interactions, with any particular individual playing a limited
role.
MLS theory can organize the study of group-level cognition in
humans, providing a framework for interpreting the existing
psychological literature and suggesting directions for future
research. For example, cooperation is required only for tasks
that exceed the capacity of individuals, yet task complexity has
seldom been manipulated as an independent variable in group-
cognition research. Using a task based on the game of 20
questions, Wilson, Timmel, & Miller (2004) showed that groups
perform better than individuals do and that the performance gap
increases with the difficulty of the word being guessed. Group
performance is uncorrelated with the performance of individual
members playing alone, and playing in a group does not sub-
sequently improve individual performance. In other words, the
advantages of playing as a group require being in a group. The
performance advantage of groups could be demonstrated even
when the task was presented in the format of a brainstorming
experiment, where advantages of real groups compared to
nominal groups (i.e., merged results of individuals playing
alone) have been notoriously difficult to demonstrate (Mullen,
Johnson, & Salas, 1991).
Although cognitive cooperation has received some attention
in psychological research (i.e., transactive memory; Wegner,
1986), MLS theory suggests that it deserves to occupy center
stage in research on group cognition and performance.
Leadership and Group Decision Making
Leadership has long occupied the attention of psychologists,
with 7,500 studies cited in the most recent Handbook of Lead-
ership. Yet, researchers frequently comment that the field is
poorly integrated. MLS theory can provide a unifying theoretical
framework for interpreting the existing literature and suggesting
new research directions (Van Vugt, 2006). Two major hypotheses
about leadership correspond directly to selection at the indi-
vidual and group levels. The first is that leadership is a by-
product of social dominance within the group. Individuals
compete for power and the winners get to make the decisions,
forcing the losers to submit to them. The second is that leader-
ship is part of an organizational structure that functions well at
the group level. Coordinating action and making collective de-
cisions for the good of the group often require leader–follower
relationships.
Making sense of human leadership requires both hypotheses.
Individual selection models suggest there are always at least
some individuals who wish to acquire power for themse lves, even
at the expense of others and the group as a whole. Without strong
mechanisms for thwarting their ambitions, leadership will take
on characteristics of dominance. Fortunately, strong mecha-
nisms for preventing exploitation do exist in human societies, as
we have seen, and they are applied with special force to leaders
so that they do not abuse the power that they have been given. For
instance, gossip and ridicule are focused primarily on important
members of a group. Domineering leaders are resented com-
pared to those who are generous, trustworthy, and empathetic. Of
course, ambition and aggression in a leader might also be assets
for the whole group, especially in competitive relationships with
other groups, which might explain why many societies have
separate leaders for war and peace (Van Vugt, 2006).
Social Identity and Intergroup Relations
Ingroup favoritism and outgroup hostility are the hallmarks of
human social psychology, and MLS theory explains why. Be-
tween-group conflict has been a major force in human history,
selecting for a range of group-level traits that may be costly for
individuals but are hugely beneficial to their group. When par-
ticular conditions are met—such as the real or imagined pres-
ence of an outgroup—these traits become apparent. Minimal
group experiments show that humans readily identify with and
discriminate against members of outgroups even if they know
that group membership is randomly decided by the flip of a coin
(Brewer, 1979). Humans are also quite prepared to make sub-
stantial sacrifices by volunteering time, donating money, or
taking risky actions to defend their group. MLS theory can
provide a theoretical framework for these well-established re-
sults in social psychology and set an agenda for future research
and practical applications (Kurzban & Neuberg, 2005).
Cultural Psychology
The human major transition enabled our ancestors to spread over
the globe—occupying hundreds of ecological niches—yet we
remained a single biological species. It is amazing that a single
species can acquire the adaptations to survive in environments
as different as the frozen arctic, the arid desert, the humid
rainforest, and remote islands thousands of miles from the
mainland. This diversification requires a fast-paced process of
cultural evolution (Richerson & Boyd, 2005), with three major
implications for psychological science.
First, cultural evolution requires a complex psychological
infrastructure that evolved by genetic evolution. An analogy with
the immune system is instructive. Our bodies are capable of
rapidly adapting to diseases, but only thanks to a genetically
evolved immune system that is mind-boggling in its complexity
and sophistication when understood in detail. Something com-
parable must exist to explain our genetically innate capacity for
8Volume 17—Number 1
Multilevel Selection Theory
rapid cultural adaptation, which should occupy center stage in
psychological research.
Second, cultural evolution can create profound psychological
differences among people, which are no less profound for being
cultural rather than genetic. Social psychologist Richard Nisbett
learned this over the course of his career, as he recounted at the
end of a recent review article:
Almost two decades ago, the senior author wrote a book with Lee
Ross entitled, modestly, Human Inference. Roy D’Andrade, a
distinguished cognitive anthropologist, read the book and pro-
nounced it a ‘‘good ethnography.’’ The author was shocked and
dismayed. But we now wholeheartedly agree with D’Andrade’s
contention about the limits of research conducted in a single
culture. Psychologists who choose not to do cross-cultural psy-
chology may have chosen to be ethnographers instead. (Nisbett,
Peng, Choi, & Norenzayan, 2001, p. 307)
Third, human activities such as music, dance, visual art,
literature, and religion, which are associated more with the
humanities than the human behavioral sciences, emerge as more
worthy of scientific study when viewed from an MLS perspective.
These activities are culturally universal (although diverse in
their specific expression), appear early in life, and—like sex—
do not require incentives to perform because they are so plea-
surable in their own right. In other words, they have all the
earmarks of genetically evolved adaptations (Baumeister, 2005).
Far from superfluous, they might play an essential role in de-
fining groups, bonding their members together, coordinating
their activities, and facilitating the social transmission of ac-
quired information.
INTEGRATING THE CONCEPT OF GROUPS AS
ORGANISMS ACROSS ALL SCIENTIFIC DISCIPLINES
It is an interesting fact, worth the attention of social historians,
that the concept of the group as an organism was widely accepted
until the middle of the 20th century, when it was rejected by
various scientific disciplines. It is now making a comeback
through the application of MLS thinking, providing a firm sci-
entific foundation for the concept of groups as organisms—not as
an axiomatic statement about all societies but as a possibility
that is realized when certain conditions are met. This foundation
can be built upon by psychology in addition to all other human-
related disciplines, in an integration of knowledge that is the
hallmark of evolutionary inquiry.
Recommended Reading
Richerson, P.J., & Boyd, R. (2005). (See References). A comprehensive
account of human cultural evolution from a multilevel evolutionary
perspective.
Wilson, D.S. & Wilson, E.O. (2007). (See References). A recent as-
sessment of multilevel selection theory for a broad academic au-
dience.
Wilson, D.S. (2007). Evolution for everyone: How Darwin’s theory can
change the way we think about our lives. New York: Delacorte Press.
The first account of evolution and its implications for human affairs
for a general audience that is based on multilevel selection theory.
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Baumeister, R.F. (2005). The cultural animal: Human nature, meaning,
and social life. New York: Oxford University Press.
Boehm, C. (1999). Hierarchy in the forest. Cambridge, MA: Harvard
University Press.
Brewer, M.B. (1979). Ingroup bias in the minimal group situation: A
cognitive-motivational analysis. Psychological Bulletin,86, 307–
324.
Campbell, D.T. (1994). How individual and face-to-face group selection
undermine firm selection in organizational evolution. In J.A.C.
Baum & J.V. Singh (Eds.), Evolutionary dynamics of organizations
(pp. 23–38). New York: Oxford University Press.
De Cremer, D., & Van Vugt, M. (1999). Social identification effects in
social dilemmas: A transformation of motives. European Journal of
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Fehr, E., & Ga
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Kurzban, R., & Neuberg, S. (2005). Managing ingroup and outgroup
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Volume 17—Number 1 9
David Sloan Wilson, Mark Van Vugt, and Rick O’Gorman
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