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Clash of the Titans

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DOI: 10.1525/bio.2012.62.11.8

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 Fall Focus on Books

Clash of the Titans

The Social Conquest of Earth. Edward E. O. Wilson defends the notion manipulation (Reeve and Keller 1997).
O. Wilson. W. W. Norton (Liveright), that humans are eusocial by recalling Therefore, he writes in Social Conquest
2012. 352 pp., illus. $27.95. (ISBN that early hominins adopted collec- on page 144,
978-0-87140-413-8 hardcover). tive child rearing in a fixed location
defended by some group members The origin and evolution
while others dispersed to forage, scav- of eusocial insects can be

E dward O. Wilson is an expert on

social insects, but early in his
career, he addressed human sociality.
enge, or hunt (Burkart et al. 2009,
Chapais 2009, Hrdy 2009, Gintis and
van Schaik 2012, Pagano 2012). All
viewed as processes driven by
individual-level natural selec-
tion. It is best tracked from
His book Sociobiology (1975) initi- known eusocial species went through queen to queen from one gen-
ated a field that has become a major a similar stage of collective brood eration to the next, with the
research area in the behavioral sci- rearing. With alloparenting (the pro- workers of each colony pro-
ences and included a controversial vision of parental care by individuals duced as phenotypic exten-
chapter on human social evolution. other than the recipient’s biological sions of the mother queen.…
In The Social Conquest of Earth, E. O. parent) in a fixed location established, The defending worker is part
Wilson updates his earlier ground- E. O. Wilson argues, there is strong of the queen’s phenotype, as
breaking treatment of Homo sapiens. selection on traits that improve coop- teeth and fingers are part of
He stakes out a strong position on erative behavior, which would culmi- your own phenotype.
the levels-of-selection debate, rejects nate in eusociality.
inclusive-fitness theory, and redefines To supporters of inclusive-fitness
eusociality to include Homo sapiens. Clash of the Titans theory, these statements are outrageous
The book is based on several articles E. O. Wilson’s theory thus diverges and strike a blow at population genet-
published in professional journals, from the widely held view that ics itself. E. O. Wilson and his coau-
including those of E. O. Wilson and human sociality can be explained by thors simply could not be believed.
Hölldobler (2005); D. S. Wilson and invoking Hamilton’s (1964a, 1964b) Because of its visibility in the journal
Wilson (2007); E. O. Wilson (2008); inclusive-fitness theory. Inclusive- Nature, Nowak, Tarnita, and Wilson’s
and, most prominently, Nowak, Tar- fitness theory shows that a person- (2010) critique of inclusive-fitness the-
nita, and Wilson (2010). ally costly behavior can spread if the ory has drawn the collective ire of a
E. O. Wilson’s work has stirred up behavior offers sufficient benefits to host of prominent population bio-
a vigorous—and sometimes highly genetically related individuals. E. O. logists. Nature subsequently published
intemperate—debate that has shaken Wilson now rejects this theory, writ- several brief communications in which
the very foundations of population ing in The Social Conquest of Earth Nowak, Tarnita, and Wilson’s (2010)
biology. In this review, I address the (p. 51) that claims were vigorously rejected, one of
fundamental issues involved and sug- which had 134 signatures (Abbot et al.
gest a resolution that may be accept- The foundations of the gen- 2011, Boomsma et al. 2011, Strass-
able to both sides of this clash of the eral theory of inclusive fitness man et al. 2011). Rousset and Lion
Titans. based on the assumptions of (2011) accused Nowak, Tarnita, and
E. O. Wilson’s treatment of euso- kin selection have crumbled, Wilson (2010) of saying nothing new
ciality in The Social Conquest of Earth while evidence for it has grown and of using “rhetorical devices.” They
is a major innovation. A species is equivocal at best.… Inclusive- then attacked the journal Nature itself,
customarily defined as eusocial if it fitness theory, often called kin arguing (p. 1390) that
is composed of colonies in each of selection theory, is both math-
which there is a reproductive divi- ematically and biologically The publication of this article
sion of labor, as in the cases of social incorrect. illustrates more general con-
bees and termites, in whose societies cerns about the publishing
a single or a few individuals (queens E. O. Wilson favors the more tradi- process.… Nature’s extravagant
and their mates) reproduce and other tional individual-fitness-maximization editorial characterization of the
castes do not. In humans, by contrast, model in understanding advanced paper as “the first mathemati-
adults are generically reproductives. eusociality, which is defined as occur- cal analysis of inclusive-fitness
ring if the workers are sterile or if theory” recklessly tramples on
worker reproduction is suppressed nearly 50 years of accumulated
doi:10.1525/bio.2012.62.11.8 by policing (Frank 1995) or queen knowledge.

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Fall Focus on Books

In passing sentence on The Social cooperate with other genes in promot- only replicators—the sole entities that
Conquest of Earth, Dawkins (2012) ing the success of the genome as a are preserved in the evolutionary pro-
concluded, “To borrow from Dorothy whole. Of course, the same argument cess. Individuals, in Dawkins’s view,
Parker, this is not a book to be tossed applies to inclusive fitness at the level are simply vehicles that genes build to
lightly aside. It should be thrown with of the individual in an advanced euso- shuttle themselves successfully from
great force.” cial colony. one body to another. Unlike genes,
By contrast, some supporters of genotypes and groups of genotypes
Hamilton’s rule and its limitations Hamilton’s rule take the notion that are transitory in sexually reproducing
Kin selection theory holds that an genes maximize inclusive fitness as the species, he argues; they are broken up
allele for helping another will spread very core of evolutionary biological and reorganized continually by sexual
if a mathematical inequality known theory. For instance, West and col- recombination.
as Hamilton’s rule holds. The rule says leagues (2011, p. 233) explained that But, in fact, genomes are replicators
that the allele will spread if the benefit “since Darwin, the only fundamen- as much as genes are, and the pheno-
from the help, in terms of fitness to tal change in our understanding of types that genomes control include
the recipient, multiplied by the recipi- adaptation has been Hamilton’s devel- metazoan individuals and social
ent’s degree of relatedness to the bene- opment of inclusive-fitness theory.… groups, such as E. O. Wilson’s eusocial
factor, is greater than the cost to the The idea [is] that organisms can be colonies and human societies. This is
benefactor. Degree of relatedness is usu- viewed as maximizing agents.” so because a gene promotes the fitness
ally understood as the probability that of its host only if it is complemented
an allele is identical in the benefactor by a well-orchestrated configura-
and the recipient by common descent. tion of genes with which it interacts
In box 1, I derive a generalization of (Sober and Lewontin 1982, Hammer-
Hamilton’s rule that relaxes key, usu- stein and Leimar 2006, Noble 2011).
ally unstated (and often implausible) Species-defining loci in the genome
assumptions about the costs and ben- are therefore necessarily homozygous
efits of helping behavior to relatives and identical in both members of a
versus nonrelatives that are essential mating pair so that crossing over and
in the derivation of the standard form meiosis do not destroy the integrity of
of Hamilton’s rule. My generalization these complementary loci. Of course,
makes clear that an allele that causes this assertion must be qualified to deal
helping can spread even if it reduces with introns and exons, dominance,
the mean fitness of the population. maternal–paternal conflicts, auto-
It follows that Hamilton’s rule does The technical literature says other- somal and sex-linked chromosomes,
not explain eusociality and, indeed, is wise. The conditions under which evolu- fitness-neutral base replacements, and
focused on what may be a minor subset tionary dynamics lead to inclusive-fitness the like. But these adjustments do not
of the biological conditions that favor maximization have been carefully stud- weaken the integrity of the genome.
the fitness of a gene and promote coop- ied by Alan Grafen and his fellow We may call the genes that support
eration in metazoan cells and social researchers, who have shown that this phenotypic commonality the spe-
species (Hammerstein and Leimar Darwinian population dynamics entail cies’ core genome. The core genome is a
2006); the absence of inclusive-fitness- inclusive-fitness maximization at the replicator and, therefore, subject to the
maximizing behavior even when Ham- individual and gene levels, but only laws of natural selection.
ilton’s rule is satisfied is widely observed assuming that fitness effects are addi-
in social species (Queller and Strass- tive (Grafen 1999, 2006, Gardner and Levels of selection
mann 1998, Nonacs 2011). The cooper- Welsh 2011, Gardner et al. 2011). But, In urging a harmonious resolution
ation of genes in metazoan cells (Leigh of course, the additivity assumption to the clash of the Titans, Okasha
1971) and in social groups (Bowles and implies that a = = 0 in the general- (2012) suggested that the level of selec-
Gintis 2011) is therefore effected by ized Hamilton rule, which is expressed tion is a matter of personal choice or
selection at a higher level than the gene in equations 3 and 5 in box 1, in ease of analysis, because mathemati-
itself. E. O. Wilson’s critique of Hamil- which case Hamilton’s rule is fully cal models show that the result is not
ton’s rule is justified. acceptable. affected by this choice: All frameworks
give the same answer. Many biologists
Genes survive because they Core genomes as replicators agree with this assessment (Queller
cooperate In his critique of The Social Conquest 1992, Dugatkin and Reeve 1994,
Genes survive the evolutionary dyna- of Earth, Dawkins (2012) repeated an Wilson DS and Sober 1994, Kerr and
mic, then, not because they maxi- argument that he first developed in Godfrey-Smith 2002, Lehmann et al.
mize inclusive fitness but because they The Selfish Gene (1976): Genes are the 2007, Wilson DS and Wilson 2007).

988 BioScience • November 2012 / Vol. 62 No. 11 www.biosciencemag.org

 Fall Focus on Books

Box 1. A generalized Hamilton’s rule.

The derivation of Hamilton’s rule is clean, intuitive, and simple. Suppose there is an allele at a locus of a haploid organism that leads
individual A to incur a fitness cost c that bestows a fitness gain b on individual B. Suppose the frequency of this allele in the population
is q, and the probability that B has a copy of the allele is p. Then, if the size of the population is n, there are qn individuals with the
helping allele, and they increase the number of members of the population from n to n + qn(b – c), but they increase the number of
helping alleles from qn to qn + qn(pb – c). Therefore, the frequency of the allele from one period to the next will increase if
qn + qn(pb – c) b(p – q)
∆q = –q= – c > 0. (1)
n + qn(b – c) 1–q

Now suppose that B has a degree of relatedness r to A, meaning that B has a copy of the helping allele with probability r by common
descent from an ancestor. Then we can write

p = r + (1 – r)q,

where the second term comes from the fact that if B does not have the helping allele by descent from a common parent, B may still
have the allele with the populationwide probability q. Substituting the above expression for p in equation 1, we arrive at Hamilton’s
rule for the allele to spread:

br > c. (2)

We can improve the realism of this equation by adding diploidy, inbreeding, dominance, mutual helping, and the like. Uyenoyama
and Feldman (1980), Michod and Hamilton (1980), Queller (1992), Rousset and Billard (2000), and many others have shown that
something resembling equation 2 will still hold, although in general the frequency q will also appear in the revised equation 2.
Suppose, however, that in performing the helping act, A induces a fitness cost on nonrelatives and a fitness cost on the colony
as a whole. For example, > 0 can occur if A redirects brooding care from nonrelative to relative larvae, and > 0 if A’s help reduces
its contribution to colonywide tasks. If the population size is n in the current period, population size n′ in the next period will include
n + qn(b – c) because of the helping behavior but will be reduced by n q and, because the number of nonrelatives is n(1 – q), will be
additionally reduced by n(1 – q) . The number of relatives of the helping allele in the current period is qn, which is increased by the
helping behavior by qn(pr – c) and decreased through lower colony efficiency by qn q. Therefore, the new population size is

n′ = n(1 – (1 – q) – q) + qn(b – c), (3)

and equation 1 becomes

qn(1 – βbq) + qn(pb – c)

∆q = – q > 0, (4)
n(1 – αq(1 – q) – βq) + qn(b – c)

which simplifies to

b(p – q) + (1 – q)q > c (1 – q).

Substituting p = r + (1 – p)q, we get what I will call the generalized Hamilton’s rule:

br + q > c. (5)

Equation 5 shows that the cost to the colony of the helping behavior ( ) is ignored in the equation for the fitness of the helping
allele and that transferring resources from nonrelatives to relatives ( > 0) can lead to an increase in the frequency of the helping allele
even if it in no other way benefits relatives (b = 0). The effect on population fitness of an increase in the helping allele is then given by
equation 6, which is derived from equation 3:

dn´
= n(b – c – α(1 – 2q) – β). (6)
dq

There is, of course, nothing to prevent this effect from being negative even when the Hamilton condition br > c is satisfied or from
being positive if even if Hamilton’s rule is violated.

However, this is true only for account- Because there are replicators at vari- to an evolutionary dynamic, selec-
ing purposes; consideration of causal ous levels of biological organization tion must also occur at various levels.
processes tells a different story. and because replicators are subject A segregation distorter, for instance,

www.biosciencemag.org November 2012 / Vol. 62 No. 11 • BioScience 989

Fall Focus on Books

operates at the level of the gene, parental reproductives can lead work- Conclusions
because selective forces operative at ers to behave in ways that reduce E. O. Wilson’s assertion that the
higher levels need not be invoked in the mean fitness of the colony, but inclusive-fitness approach is “both
its explanation (Sober and Lewontin such behavior is not an expression mathematically and biologically incor-
1982, Burt and Trivers 2006). Where of the workers’ inclusive fitness but, rect” should be revised. A more accu-
the core genome specifies structured rather, a transcription error. Such rate assessment is that Hamilton’s rule
interactions among related individu- “outlaw” workers are not “policed” is incomplete, and with a few changes,
als, kin selection occurs and is there- but are, rather, eliminated or con- it can maintain its exalted position
fore a form of group selection. trolled through an error correction in sociobiology. These changes lead
Where the causal relations inscribed process within the colony. Like cogs in us to the generalized Hamilton’s rule,
in the core genome occur between or a machine, a worker in such a colony which points beyond itself to the need
among metazoan individuals, analysis cannot “defect,” although it can “fail” for many additional tools in the biolo-
in terms of group selection at a level as a part in a machine (Hammerstein gist’s repertoire to explain complex
higher than the individual is appro- and Leimar 2006). Richard Dawkins social behavior.
priate. The group could merely affect could have explained all this in his The evolutionary forces that deter-
fitness payoffs to its members, but famous book The Extended Phenotype mine the nonlinear interactions
generally, this will entail that group (1989); he might have saved a lot of among loci in metazoans and among
structure be inscribed in the core futile debate. individuals in sociobiological species
genome. Social groups, then, are sub- must be studied using tools such as
ject to exactly the same evolution- Homo sapiens is not a eusocial the phenotypic gambit (Grafen 1984),
ary forces—replication with mutation, species game theory (DS Wilson 1977, 1980,
selection, and heredity—as are indi- Humans evolved in cohesive groups Taylor 1992, 1996), agent-based mod-
vidual genes and metazoan individuals. involving many unrelated families eling (Gintis 2009), and the physiol-
Whatever the level at which selection (Boyd and Silk 2002) and lack a repro- ogy of suppressor genes (Leigh 1977).
operates, the resulting dynamic can, of ductive division of labor (but see Foster E. O. Wilson and the coauthors of his
course, at least in principle, be trans- and Ratnieks 2005). Moreover, human recent publications are to be lauded for
lated into changes in gene frequencies. society sustains a high level of coop- expanding our horizons, not scorned
eration among genealogically distant for their small mistakes and tendency
No altruism in advanced eusocial individuals (Gintis 2000, Richerson to overstate their position. So The
species and Boyd 2004, Bowles and Gintis Social Conquest of Earth is, despite
Biological altruism is defined as a 2011). By contrast, the reproductive some weaknesses, a book that I recom-
behavior that lowers the personal fit- division of labor is central to sustain- mend. It is an inspiring account of a
ness of the focal individual but raises ing cooperation in eusocial species, significant advance in our understand-
the fitness of other individuals (Foster just as sequestration of the germline ing. Contemporary inclusive-fitness
2008). In the case of advanced euso- is key to the emergence of complex theory is a very big boat, and it can
ciality, worker fitness is zero (i.e., c = 0 metazoans (Buss 1987). Thus I argue, only be rocked by giants (Gadagkar
in equations 2 and 5 in box 1). Altru- contra E. O. Wilson, that it is prudent 2010).
ism in advanced eusocial species is to exclude humans from the eusocial
therefore precluded in principle. Even fold. Acknowledgments
the generalized Hamilton’s rule cannot Much of E. O. Wilson’s critique of I would like to thank Samuel Bowles,
explain altruism in advanced eusocial the inclusive-fitness model of eusocial- Steven Frank, and David Queller for
species, because it does not exist. ity does not apply directly to humans advice in preparing this article.
In an advanced eusocial colony, or to primitively social species. In our
E. O. Wilson maintains, the workers work, my colleagues and I routinely References cited
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queens and males than would be pos- Griffin AS, Holman L, Hughes WOH, Keller
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Herbert Gintis ([email protected])
Forthcoming. Queller DC, Strassmann JE. 1998. Kin selection
is a professor of economics at Central
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and group selection. Pages 62–84 in Krebs Reeve HK, Keller L. 1997. Reproductive bribing European University, in Budapest; and
JR, Davies NB, eds. Behavioural Ecology: An and policing as evolutionary mechanisms a visiting professor at the University
Evolutionary Approach Sinauer. for the suppression of within-group selfish- of Siena, in Siena, Italy. He is also
———. 1999. Formal Darwinism, the individual- ness. American Naturalist 150 (suppl. 1): an external professor at the Santa Fe
as-maximizing-agent analogy and bet-hedging. S42–S58. Institute, in Santa Fe, New Mexico.

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