Biological Altruism

First published Tue Jun 3, 2003; substantive revision Sun Jul 21, 2013
In evolutionary biology, an organism is said to behave altruistically when its behaviour benefits
other organisms, at a cost to itself. The costs and benefits are measured in terms of reproductive
fitness, or expected number of offspring. So by behaving altruistically, an organism reduces the
number of offspring it is likely to produce itself, but boosts the number that other organisms are
likely to produce. This biological notion of altruism is not identical to the everyday concept. In
everyday parlance, an action would only be called altruistic if it was done with the conscious
intention of helping another. But in the biological sense there is no such requirement. Indeed, some
of the most interesting examples of biological altruism are found among creatures that are
(presumably) not capable of conscious thought at all, e.g. insects. For the biologist, it is the
consequences of an action for reproductive fitness that determine whether the action counts as
altruistic, not the intentions, if any, with which the action is performed.
Altruistic behaviour is common throughout the animal kingdom, particularly in species with
complex social structures. For example, vampire bats regularly regurgitate blood and donate it to
other members of their group who have failed to feed that night, ensuring they do not starve. In
numerous bird species, a breeding pair receives help in raising its young from other helper birds,
who protect the nest from predators and help to feed the fledglings. Vervet monkeys give alarm
calls to warn fellow monkeys of the presence of predators, even though in doing so they attract
attention to themselves, increasing their personal chance of being attacked. In social insect colonies
(ants, wasps, bees and termites), sterile workers devote their whole lives to caring for the queen,
constructing and protecting the nest, foraging for food, and tending the larvae. Such behaviour is
maximally altruistic: sterile workers obviously do not leave any offspring of their ownso have
personal fitness of zerobut their actions greatly assist the reproductive efforts of the queen.
From a Darwinian viewpoint, the existence of altruism in nature is at first sight puzzling, as Darwin
himself realized. Natural selection leads us to expect animals to behave in ways that increase their
own chances of survival and reproduction, not those of others. But by behaving altruistically an
animal reduces its own fitness, so should be at a selective disadvantage vis--vis one which behaves
selfishly. To see this, imagine that some members of a group of Vervet monkeys give alarm calls
when they see predators, but others do not. Other things being equal, the latter will have an
advantage. By selfishly refusing to give an alarm call, a monkey can reduce the chance that it will
itself be attacked, while at the same time benefiting from the alarm calls of others. So we should
expect natural selection to favour those monkeys that do not give alarm calls over those that do. But
this raises an immediate puzzle. How did the alarm-calling behaviour evolve in the first place, and
why has it not been eliminated by natural selection? How can the existence of altruism be
reconciled with basic Darwinian principles?
1. Altruism and the Levels of Selection
2. Kin Selection and Inclusive Fitness
2.1 A Simple Illustration: the Prisoner's dilemma
3. Conceptual Issues
3.1 Altruism, Co-operation, Mutualism
3.2 Weak and Strong Altruism
3.3 Short-term versus Long-term Fitness Consequences
4. Reciprocal Altruism
5. But is it Real Altruism?
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1. Altruism and the Levels of Selection

The problem of altruism is intimately connected with questions about the level at which natural
selection acts. If selection acts exclusively at the individual level, favouring some individual
organisms over others, then it seems that altruism cannot evolve, for behaving altruistically is
disadvantageous for the individual organism itself, by definition. However, it is possible that
altruism may be advantageous at the group level. A group containing lots of altruists, each ready to
subordinate their own selfish interests for the greater good of the group, may well have a survival
advantage over a group composed mainly or exclusively of selfish organisms. A process of
between-group selection may thus allow the altruistic behaviour to evolve. Within each group,
altruists will be at a selective disadvantage relative to their selfish colleagues, but the fitness of the
group as a whole will be enhanced by the presence of altruists. Groups composed only or mainly of
selfish organisms go extinct, leaving behind groups containing altruists. In the example of the
Vervet monkeys, a group containing a high proportion of alarm-calling monkeys will have a
survival advantage over a group containing a lower proportion. So conceivably, the alarm-calling
behaviour may evolve by between-group selection, even though within each group, selection
favours monkeys that do not give alarm calls.
The idea that group selection might explain the evolution of altruism was first broached by Darwin
himself. In The Descent of Man (1871), Darwin discussed the origin of altruistic and self-sacrificial
behaviour among humans. Such behaviour is obviously disadvantageous at the individual level, as
Darwin realized: he who was ready to sacrifice his life, as many a savage has been, rather than
betray his comrades, would often leave no offspring to inherit his noble nature (p.163). Darwin
then argued that self-sacrificial behaviour, though disadvantageous for the individual savage,
might be beneficial at the group level: a tribe including many members who...were always ready to
give aid to each other and sacrifice themselves for the common good, would be victorious over
most other tribes; and this would be natural selection (p.166). Darwin's suggestion is that the
altruistic behaviour in question may have evolved by a process of between-group selection.
The concept of group selection has a chequered and controversial history in evolutionary biology.
The founders of modern neo-DarwinismR.A. Fisher, J.B.S. Haldane and S. Wrightwere all
aware that group selection could in principle permit altruistic behaviours to evolve, but they
doubted the importance of this evolutionary mechanism. Nonetheless, many mid-twentieth century
ecologists and some ethologists, notably Konrad Lorenz, routinely assumed that natural selection
would produce outcomes beneficial for the whole group or species, often without even realizing that
individual-level selection guarantees no such thing. This uncritical good of the species tradition
came to an abrupt halt in the 1960s, due largely to the work of G.C. Williams (1966) and J.
Maynard Smith (1964). These authors argued that group selection was an inherently weak
evolutionary force, hence unlikely to promote interesting altruistic behaviours. This conclusion was
supported by a number of mathematical models, which apparently showed that group selection
would only have significant effects for a limited range of parameter values. As a result, the notion
of group selection fell into widespread disrepute in orthodox evolutionary circles; see Sober and
Wilson 1998, Segestrale 2000, Okasha 2006, Leigh 2010 and Sober 2011 for details of the history
of this debate.
The major weakness of group selection as an explanation of altruism, according to the consensus
that emerged in the 1960s, was a problem that Dawkins (1976) called subversion from within; see
also Maynard Smith 1964. Even if altruism is advantageous at the group level, within any group
altruists are liable to be exploited by selfish free-riders who refrain from behaving altruistically.
These free-riders will have an obvious fitness advantage: they benefit from the altruism of others,
but do not incur any of the costs. So even if a group is composed exclusively of altruists, all
behaving nicely towards each other, it only takes a single selfish mutant to bring an end to this
happy idyll. By virtue of its relative fitness advantage within the group, the selfish mutant will out-
reproduce the altruists, hence selfishness will eventually swamp altruism. Since the generation time
of individual organisms is likely to be much shorter than that of groups, the probability that a selfish
mutant will arise and spread is very high, according to this line of argument. Subversion from
within is generally regarded as a major stumbling block for group-selectionist theories of the
evolution of altruism.
If group selection is not the correct explanation for how the altruistic behaviours found in nature
evolved, then what is? In the 1960s and 1970s a rival theory emerged: kin selection or inclusive
fitness theory, due originally to Hamilton (1964). This theory, discussed in detail below, apparently
showed how altruistic behaviour could evolve without the need for group-level selection, and
quickly gained prominence among biologists interested in the evolution of social behaviour; the
empirical success of kin selection theory contributed to the demise of the group selection concept.
However, the precise relation between kin and group selection is a source of ongoing controversy
(see for example the recent exchange in Nature between Nowak, Tarnita and Wilson 2010 and
Abbot et. al. 2011). Since the 1990s, proponents of multi-level selection theory have resuscitated a
form of group-level selectionsometimes called new group selectionand shown that it can
permit altruism to evolve (cf. Sober and Wilson 1998). But new group selection turns out to be
mathematically equivalent to kin selection in most if not all cases, as a number of authors have
emphasized (Grafen 1984, Frank 1998, West et al. 2007, Lehmann et al. 2007, Marshall 2011); this
point was already appreciated by Hamilton (1975). Since the relation between old and new group
selection is itself a point of controversy, this explains why disagreement about the relation between
kin and group selection should persist.

2. Kin Selection and Inclusive Fitness

The basic idea of kin selection is simple. Imagine a gene which causes its bearer to behave
altruistically towards other organisms, e.g. by sharing food with them. Organisms without the gene
are selfishthey keep all their food for themselves, and sometimes get handouts from the altruists.
Clearly the altruists will be at a fitness disadvantage, so we should expect the altruistic gene to be
eliminated from the population. However, suppose that altruists are discriminating in who they
share food with. They do not share with just anybody, but only with their relatives. This
immediately changes things. For relatives are genetically similarthey share genes with one
another. So when an organism carrying the altruistic gene shares his food, there is a certain
probability that the recipients of the food will also carry copies of that gene. (How probable
depends on how closely related they are.) This means that the altruistic gene can in principle spread
by natural selection. The gene causes an organism to behave in a way which reduces its own fitness
but boosts the fitness of its relativeswho have a greater than average chance of carrying the gene
themselves. So the overall effect of the behaviour may be to increase the number of copies of the
altruistic gene found in the next generation, and thus the incidence of the altruistic behaviour itself.
Though this argument was hinted at by Haldane in the 1930s, and to a lesser extent by Darwin in his
discussion of sterile insect castes in The Origin of Species, it was first made explicit by William
Hamilton (1964) in a pair of seminal papers. Hamilton demonstrated rigorously that an altruistic
gene will be favoured by natural selection when a certain condition, known as Hamilton's rule, is
satisfied. In its simplest version, the rule states that b > c/r, where c is the cost incurred by the
altruist (the donor), b is the benefit received by the recipients of the altruism, and r is the co-
efficient of relationship between donor and recipient. The costs and benefits are measured in terms
of reproductive fitness. The co-efficient of relationship depends on the genealogical relation
between donor and recipientit is defined as the probability that donor and recipient share genes at
a given locus that are identical by descent. (Two genes are identical by descent if they are copies
of a single gene in a shared ancestor.) In a sexually reproducing diploid species, the value of r for
full siblings is , for parents and offspring , for grandparents and grandoffspring , for full
cousins 1/8, and so-on. The higher the value of r, the greater the probability that the recipient of the
altruistic behaviour will also possess the gene for altruism. So what Hamilton's rule tells us is that a
gene for altruism can spread by natural selection, so long as the cost incurred by the altruist is offset
by a sufficient amount of benefit to sufficiently closed related relatives. The proof of Hamilton's
rule relies on certain non-trivial assumptions; see Frank 1998, Grafen 1985, 2006, Queller 1992a,
1992b, Boyd and McIlreath 2006 and Birch forthcoming for details.
Though Hamilton himself did not use the term, his idea quickly became known as kin selection,
for obvious reasons. Kin selection theory predicts that animals are more likely to behave
altruistically towards their relatives than towards unrelated members of their species. Moreover, it
predicts that the degree of altruism will be greater, the closer the relationship. In the years since
Hamilton's theory was devised, these predictions have been amply confirmed by empirical work.
For example, in various bird species, it has been found that helper birds are much more likely to
help relatives raise their young, than they are to help unrelated breeding pairs. Similarly, studies of
Japanese macaques have shown that altruistic actions, such as defending others from attack, tend to
be preferentially directed towards close kin. In most social insect species, a peculiarity of the
genetic system known as haplodiploidy means that females on average share more genes with
their sisters than with their own offspring. So a female may well be able to get more genes into the
next generation by helping the queen reproduce, hence increasing the number of sisters she will
have, rather than by having offspring of her own. Kin selection theory therefore provides a neat
explanation of how sterility in the social insects may have evolved by Darwinian means. (Note,
however, that the precise significance of haplodiploidy for the evolution of worker sterility is a
controversial question; see Maynard Smith and Szathmary 1995 ch.16, Gardner, Alpedrinha and
West 2012.)
Kin selection theory is often presented as a triumph of the gene's-eye view of evolution, which
sees organic evolution as the result of competition among genes for increased representation in the
gene-pool, and individual organisms as mere vehicles that genes have constructed to aid their
propagation (Dawkins 1976, 1982). The gene's eye-view is certainly the easiest way of
understanding kin selection, and was employed by Hamilton himself in his 1964 papers. Altruism
seems anomalous from the individual organism's point of view, but from the gene's point of view it
makes good sense. A gene wants to maximize the number of copies of itself that are found in the
next generation; one way of doing that is to cause its host organism to behave altruistically towards
other bearers of the gene, so long as the costs and benefits satisfy the Hamilton inequality. But
interestingly, Hamilton showed that kin selection can also be understood from the organism's point
of view. Though an altruistic behaviour which spreads by kin selection reduces the organism's
personal fitness (by definition), it increases what Hamilton called the organism's inclusive fitness.
An organism's inclusive fitness is defined as its personal fitness, plus the sum of its weighted effects
on the fitness of every other organism in the population, the weights determined by the coefficient
of relationship r. Given this definition, natural selection will act to maximise the inclusive fitness of
individuals in the population (Grafen 2006). Instead of thinking in terms of selfish genes trying to
maximize their future representation in the gene-pool, we can think in terms of organisms trying to
maximize their inclusive fitness. Most people find the gene's eye approach to kin selection
heuristically simpler than the inclusive fitness approach, but mathematically they are in fact
equivalent (Michod 1982, Frank 1998, Boyd and McIlreath 2006, Grafen 2006).
Contrary to what is sometimes thought, kin selection does not require that animals must have the
ability to discriminate relatives from non-relatives, less still to calculate coefficients of relationship.
Many animals can in fact recognize their kin, often by smell, but kin selection can operate in the
absence of such an ability. Hamilton's inequality can be satisfied so long as an animal behaves
altruistically towards others animals that are in fact its relatives. The animal might achieve this by
having the ability to tell relatives from non-relatives, but this is not the only possibility. An
alternative is to use some proximal indicator of kinship. For example, if an animal behaves
altruistically towards those in its immediate vicinity, then the recipients of the altruism are likely to
be relatives, given that relatives tend to live near each other. No ability to recognize kin is
presupposed. Cuckoos exploit precisely this fact, free-riding on the innate tendency of birds to care
for the young in their nests.
Another popular misconception is that kin selection theory is committed to genetic determinism,
the idea that genes rigidly determine or control behaviour. Though some sociobiologists have made
incautious remarks to this effect, evolutionary theories of behaviour, including kin selection, are not
committed to it. So long as the behaviours in question have a genetical component, i.e. are
influenced to some extent by one or more genetic factor, then the theories can apply. When
Hamilton (1964) talks about a gene which causes altruism, this is really shorthand for a gene
which increases the probability that its bearer will behave altruistically, to some degree. This is
much weaker than saying that the behaviour is genetically determined, and is quite compatible
with the existence of strong environmental influences on the behaviour's expression. Kin selection
theory does not deny the truism that all traits are affected by both genes and environment. Nor does
it deny that many interesting animal behaviours are transmitted through non-genetical means, such
as imitation and social learning (Avital and Jablonka 2000).
The importance of kinship for the evolution of altruism is very widely accepted today, on both
theoretical and empirical grounds. However, kinship is really only a way of ensuring that altruists
and recipients both carry copies of the altruistic gene, which is the fundamental requirement. If
altruism is to evolve, it must be the case that the recipients of altruistic actions have a greater than
average probability of being altruists themselves. Kin-directed altruism is the most obvious way of
satisfying this condition, but there are other possibilities too (Hamilton 1975, Sober and Wilson
1998, Bowles and Gintis 2011, Gardner and West 2011). For example, if the gene that causes
altruism also causes animals to favour a particular feeding ground (for whatever reason), then the
required correlation between donor and recipient may be generated. It is this correlation, however
brought about, that is necessary for altruism to evolve. This point was noted by Hamilton himself in
the 1970s: he stressed that the coefficient of relationship of his 1964 papers should really be
replaced with a more general correlation coefficient, which reflects the probability that altruist and
recipient share genes, whether because of kinship or not (Hamilton 1970, 1972, 1975). This point is
theoretically important, and has not always been recognized; but in practice, kinship remains the
most important source of statistical associations between altruists and recipients (Maynard Smith
1998, Okasha 2002, West et al. 2007).

2.1 A Simple Illustration: the Prisoner's dilemma

The fact that correlation between donor and recipient is the key to the evolution of altruism can be
illustrated via a simple one shot Prisoner's dilemma game. Consider a large population of
organisms who engage in a social interaction in pairs; the interaction affects their biological fitness.
Organisms are of two types: selfish (S) and altruistic (A). The latter engage in pro-social behaviour,
thus benefiting their partner but at a cost to themselves; the former do not. So in a mixed (S,A) pair,
the selfish organism does betterhe benefits from his partner's altruism without incurring any cost.
However, (A,A) pairs do better than (S,S) pairsfor the former work as a co-operative unit, while
the latter do not. The interaction thus has the form of a one-shot Prisoner's dilemma, familiar from
game theory. Illustrative payoff values to each player, i.e., each partner in the interaction,
measured in units of biological fitness, are shown in the matrix below.
Player 2
Altruist Selfish
 Altruist 11,11 0,20
Player 1
Selfish 20,0 5,5
Payoffs for (Player 1, Player 2) in units of reproductive fitness

The question we are interested in is: which type will be favoured by selection? To make the analysis
tractable, we make two simplifying assumptions: that reproduction is asexual, and that type is
perfectly inherited, i.e., selfish (altruistic) organisms give rise to selfish (altruistic) offspring.
Modulo these assumptions, the evolutionary dynamics can be determined very easily, simply by
seeing whether the S or the A type has higher fitness, in the overall population. The fitness of the S
type, W(S), is the weighted average of the payoff to an S when partnered with an S and the payoff to
an S when partnered with an A, where the weights are determined by the probability of having the
partner in question. Therefore,
W(S) = 5 * Prob(S partner/S) + 20 * Prob(A partner/S)

(The conditional probabilities in the above expression should be read as the probability of having a
selfish (altruistic) partner, given that one is selfish oneself.)
Similarly, the fitness of the A type is:
W(A) = 0 * Prob(S partner/A) + 11 * Prob(A partner/A)

From these expressions for the fitnesses of the two types of organism, we can immediately deduce
that the altruistic type will only be favoured by selection if there is a statistical correlation between
partners, i.e., if altruists have greater than random chance of being paired with other altruists, and
similarly for selfish types. For suppose there is no such correlationas would be the case if the
pairs were formed by random sampling from the population. Then, the probability of having a
selfish partner would be the same for both S and A types, i.e., P(S partner/S) = P(S partner/A).
Similarly, P(A partner/S) = P(A partner/A). From these probabilistic equalities, it follows
immediately that W(S) is greater than W(A), as can be seen from the expressions for W(S) and W(A)
above; so the selfish type will be favoured by natural selection, and will increase in frequency every
generation until all the altruists are eliminated from the population. Therefore, in the absence of
correlation between partners, selfishness must win out (cf. Skyrms 1996). This confirms the point
noted in section 2that altruism can only evolve if there is a statistical tendency for the
beneficiaries of altruistic actions to be altruists themselves.
If the correlation between partners is sufficiently strong, in this simple model, then it is possible for
the condition W(A) > W(S) to be satisfied, and thus for altruism to evolve. The easiest way to see
this is to suppose that the correlation is perfect, i.e., selfish types are always paired with other
selfish types, and ditto for altruists, so P(S partner/S) = P(A partner/A) = 1. This assumption implies
that W(A)=11 and W(S)=5, so altruism evolves. With intermediate degrees of correlation, it is also
possible for the condition W(S) > W(A) to be satisfied, given the particular choice of payoff values
in the model above.
This simple model also highlights the point made previously, that donor-recipient correlation, rather
than genetic relatedness, is the key to the evolution of altruism. What is needed for altruism to
evolve, in the model above, is for the probability of having a partner of the same type as oneself to
be sufficiently larger than the probability of having a partner of opposite type; this ensures that the
recipients of altruism have a greater than random chance of being fellow altruists, i.e., donor-
recipient correlation. Whether this correlation arises because partners tend to be relatives, or
because altruists are able to seek out other altruists and choose them as partners, or for some other
reason, makes no difference to the evolutionary dynamics, at least in this simple example.
3. Conceptual Issues
Altruism is a well understood topic in evolutionary biology; the theoretical ideas explained above
have been extensively analysed, empirically confirmed, and are widely accepted. Nonetheless, there
are a number of conceptual ambiguities surrounding altruism and related concepts in the literature;
some of these are purely semantic, others are more substantive. Three such ambiguities are briefly
discussed below; for further discussion, see West et al. 2007, Sachs et al. 2004 or Lehmann and
Keller 2006.

3.1 Altruism, Co-operation, Mutualism

According to the standard definition, a social behaviour counts as altruistic if it reduces the fitness
of the organism performing the behaviour, but boosts the fitness of others. This was the definition
used by Hamilton (1964), and by many subsequent authors. However, there is less consensus on
how to describe behaviours that boost the fitness of others but also boost the fitness of the organism
performing the behaviour. As West et al. (2007) note, such behaviours are sometimes termed co-
operative, but this usage is not universal; others use co-operation to refer to behaviour that boosts
the fitness of others irrespective of its effect on self; while still others use cooperation as a
synonym for altruism. (Indeed, in the simple Prisoner's dilemma game above, the two strategies are
usually called co-operate and defect.) To avoid this confusion, West et al. (2007) suggest the
term mutual benefit for behaviours that benefit both self and other, while Sachs et al. (2004)
suggest byproduct benefit.
Whatever term is used, the important point is that behaviours that benefit both self and others can
evolve much more easily than altruistic behaviours, and thus require no special mechanisms such as
kinship. The reason is clear: organisms performing such behaviours thereby increase their personal
fitness, so are at a selective advantage vis-a-vis those not performing the behaviour. The fact that
the behaviour has a beneficial effect on the fitness of others is a mere side-effect, or byproduct, and
is not part of the explanation for why the behaviour evolves. For example, Sachs et al. (2004) note
that an action such as joining a herd or a flock may be of this sort; the individual gains directly, via
his reduced risk of predation, while simultaneously reducing the predation risk of other individuals.
By contrast with an altruistic action, there is no personal incentive to cheat, i.e., to refrain from
performing the action, for doing so would directly reduce personal fitness.
Also indicative of the difference between altruistic behaviour and behaviour that benefit both self
and others is the fact that in the latter case, though not the former, the beneficiary may be a member
of a different species, without altering the evolutionary dynamics of the behaviour. Indeed, there are
numerous examples where the self-interested activities of one organism produce an incidental
benefit for a non-conspecific; such behaviours are sometimes called mutualistic, though again, this
is not the only way that the latter term has been used (West et al. 2007). By contrast, in the case of
altruism, it makes an enormous difference whether the beneficiary and the donor are con-specifics
or not; for if not, then kin selection can play no role, and it is quite unclear how the altruistic
behaviour can evolve. Unsurprisingly, virtually all the bona fide examples of biological altruism in
the living world involve donors and recipients that are con-specifics. (Cases of so-called reciprocal
altruism are sometimes thought to be exceptions to this generalization; but see section 4 below.)

3.2 Weak and Strong Altruism

A quite different ambiguity concerns the distinction between weak and strong altruism, in the
terminology of D.S. Wilson (1977, 1980, 1990). This distinction is about whether the altruistic
action entails an absolute or relative fitness reduction for the donor. To count as strongly altruistic, a
behaviour must reduce the absolute fitness (i.e., number of offspring) of the donor. Strong altruism
is the standard notion of altruism in the literature, and was assumed above. To count as weakly
altruistic, an action need only reduce the relative fitness of the donor, i.e., its fitness relative to that
of the recipient. Thus for example, an action which causes an organism to leave an additional 10
offspring, but causes each organism(s) with which it interacts to leave an additional 20 offspring, is
weakly but not strongly altruistic. The action boosts the absolute fitness of the donor, but boosts
the absolute fitness of other organisms by even more, thus reducing the donor's relative fitness.
Should weakly altruistic behaviours be classified as altruistic or selfish? This question is not merely
semantic; for the real issue is whether the conditions under which weak altruism can evolve are
relevantly similar to the conditions under which strong altruism can evolve, or not. Many authors
argue that the answer is no, on the grounds that weakly altruistic behaviours are individually
advantageous, so can evolve with no component of kin selection or donor-recipient correlation,
unlike strongly altruistic behaviours (Grafen 1984, Nunney 1985, West et al. 2007). To appreciate
this argument, consider a game-theoretic scenario similar to the one-shot Prisoner's dilemma of
section 4, in which organisms engage in a pair-wise interaction that affects their fitness. Organisms
are of two types, weakly altruistic (W) and non-altruistic (N). W-types perform an action that boosts
their own fitness by 10 units and the fitness of their partner by 20 units; N-types do not perform the
action. The payoff matrix is thus:
Player 2
Weak Altruist Non
Weak Altruist 30,30 10,20
Player 1
Non 20,10 0,0
Payoffs for (Player 1, Player 2) in units of reproductive fitness

The payoff matrix highlights the fact that weak altruism is individually advantageous, and thus the
oddity of thinking of it it as altruistic rather than selfish. To see this, assume for a moment that the
game is being played by two rational agents, as in classical game theory. Clearly, the rational
strategy for each individual is W, for W dominates N. Each individual gets a higher payoff from
playing W than N, irrespective of what its opponent does30 rather than 20 if the opponent plays
W, 10 rather than 0 if the opponent plays N. This captures a clear sense in which weak altruism is
individually advantageous.
In the context of evolutionary game theory, where the game is being played by pairs of organisms
with hard-wired strategies, the counterpart of the fact that W dominates N is the fact that W can
spread in the population even if pairs are formed at random (cf. Wilson 1980). To see this, consider
the expressions for the overall population-wide fitnesses of W and N:
W(W) = 30 * Prob(W partner/W) + 10 * Prob(N partner/W)

W(N) = 20 * Prob(W partner/N) + 0 * Prob(N partner/N)

(As before, Prob(W partner/W) denotes the conditional probability of having a weakly altruistic
partner given that one is weakly altruistic oneself, and so-on.) From these expressions, it is easy to
see that W(W) > W(N) even if the there is no correlation among partners, i.e., even if Prob(W
partner/W) = P(W partner/N) and P(N partner/W) = P(N partner/N). Therefore, weak altruism can
evolve in the absence of donor-recipient correlation; as we saw, this is not true of strong altruism.
So weak and strong altruism evolve by different evolutionary mechanisms, hence should not be co-
classified, according to this argument.
However, there is a counter argument due to D.S. Wilson (1977, 1980), who maintains that weak
altruism cannot evolve by individual selection alone; a component of group selection is needed.
Wilson's argument stems from the fact that in a mixed (W,N) pair, the non-altruist is fitter than the
weak altruist. More generally, within a single group of any size containing weak altruists and non-
altruists, the latter will be fitter. So weak altruism can only evolve, Wilson argues, in a multi-group
settingin which the within-group selection in favour of N, is counteracted by between-group
selection in favour of W. (On Wilson's view, the evolutionary game described above is a multi-group
setting, involving a large number of groups of size two.) Thus weak altruism, like strong altruism,
in fact evolves because it is group-advantageous, Wilson argues.
The dispute between those who regard weak altruism as individually advantageous, and those like
Wilson who regard it as group advantageous, stems ultimately from differing conceptions of
individual and group selection. For Wilson, individual selection means within-group selection, so to
determine which strategy is favoured by individual selection, one must compare the fitnesses of W
and N types within a group, or pair. For other theorists, individual selection means selection based
on differences in individual phenotype, rather than social context; so to determine which strategy is
favoured by individual selection, one must compare the fitnesses of W and N types in the same
social context, i.e., with the same partner. These two comparisons yield different answers to the
question of whether weak altruism is individually advantageous. Thus the debate over how to
classify weak altruism is intimately connected to the broader levels of selection question; see
Nunney 1985, Okasha 2005, 2006, Fletcher and Doebeli 2006, West et al. 2007, for further
discussion.

3.3 Short-term versus Long-term Fitness Consequences

A further source of ambiguity in the definition of biological altruism concerns the time-scale over
which fitness is measured. Conceivably, an animal might engage in a social behaviour which
benefits another and reduces its own (absolute) fitness in the short-term; however, in the long-term,
the behaviour might be to the animal's advantage. So if we focus on short-term fitness effects, the
behaviour will seem altruistic; but if we focus on lifetime fitness, the behaviour will seem selfish
the animal's lifetime fitness would be reduced if it did not perform the behaviour.
Why might a social behaviour reduce an animal's short-term fitness but boost its lifetime fitness?
This could arise in cases of directed reciprocation, where the beneficiary of the behaviour returns
the favour at some point in the future (cf. Sachs et al. 2004). By performing the behaviour, and
suffering the short-term cost, the animal thus ensures (or raises the chance) that it will receive return
benefits in the future. Similarly, in symbioses between members of different species, it may pay an
organism to sacrifice resources for the benefit of a symbiont with which it has a long-term
relationship, as its long-term welfare may be heavily dependent on the symbiont's welfare.
From a theoretical point of view, the most satisfactory resolution of this ambiguity is to use lifetime
fitness as the relevant parameter (cf. West et al. 2007) Thus an action only counts as altruistic if it
reduces an organism's lifetime fitness. This stipulation makes sense, since it preserves the key idea
that the evolution of altruism requires statistical association between donor and recipient; this would
not be true if short-term fitness were used to define altruism, for behaviours which reduce short-
term fitness but boost lifetime fitness can evolve with no component of kin selection, or donor-
recipient correlation. However, the stipulation has two disadvantages: (i) it makes it harder to tell
whether a given behaviour is altruistic, since lifetime fitness is notoriously difficult to estimate; (ii)
it has the consequence that most models of reciprocal altruism are mis-named.

4. Reciprocal Altruism
The theory of reciprocal altruism was originally developed by Trivers (1971), as an attempt to
explain cases of (apparent) altruism among unrelated organisms, including members of different
species. (Clearly, kin selection cannot help explain altruism among non-relatives.) Trivers' basic
idea was straightforward: it may pay an organism to help another, if there is an expectation of the
favour being returned in the future. (If you scratch my back, I'll scratch yours.) The cost of helping
is offset by the likelihood of the return benefit, permitting the behaviour to evolve by natural
selection. Trivers termed with evolutionary mechanism reciprocal altruism.
For reciprocal altruism to work, there is no need for the two individuals to be relatives, nor even to
be members of the same species. However, it is necessary that individuals should interact with each
more than once, and have the ability to recognize other individuals with whom they have interacted
in the past.[1] If individuals interact only once in their lifetimes and never meet again, there is
obviously no possibility of return benefit, so there is nothing to be gained by helping another.
However, if individuals encounter each other frequently, and are capable of identifying and
punishing cheaters who have refused to help in the past, then the helping behaviour can evolve. A
cheat who refuses to help will ultimately sabotage his own interests, for although he does not incur
the cost of helping others, he forfeits the return benefits tooothers will not help him in the future.
This evolutionary mechanism is most likely to work where animals live in relatively small groups,
increasing the likelihood of multiple encounters.
As West et al. (2007) and Bowles and Gintis (2011) note, if altruism is defined by reference to
lifetime fitness, then Trivers' theory is not really about the evolution of altruism at all; for
behaviours that evolve via reciprocation of benefits, as described by Trivers, are ultimately of direct
benefit to the individuals performing them, so do not reduce lifetime fitness. Despite this
consideration, the label reciprocal altruism is well-entrenched in the literature, and the
evolutionary mechanism that it describes is of some importance, whatever it is called. Where
reciprocal altruism is referred to below, it should be remembered that the behaviours in question are
only altruistic in the short-term.
The concept of reciprocal altruism is closely related to the Tit-for-Tat strategy in the iterated
Prisoner's Dilemma (IPD) from game theory. In the IPD, players interact on multiple occasions, and
are able to adjust their behaviour depending on what their opponent has done in previous rounds.
There are two possible strategies, co-operate and defect; the payoff matrix (per interaction) is as in
section 2.1 above. The fact that the game is iterated rather than one-shot obviously changes the
optimal course of action; defecting is no longer necessarily the best option, so long as the
probability of subsequent encounters is sufficiently high. In their famous computer tournament in
which a large number of strategies were pitted against each other in the IPD, Axelrod and Hamilton
(1981) found that the Tit-for-Tat strategy yielded the highest payoff. In Tit-For-Tat, a player follows
two basic rules: (i) on the first encounter, cooperate; (ii) on subsequent encounters, do what your
opponent did on the previous encounter. The success of Tit-for-Tat was widely taken to confirm the
idea that with multiple encounters, natural selection could favour social behaviours that entail a
short-term fitness cost. Subsequent work in evolutionary game theory, much of it inspired by
Axelrod and Hamilton's ideas, has confirmed that repeated games permit the evolution of social
behaviours that cannot evolve in one-shot situations (cf. Nowak 2006); this is closely related to the
so-called 'folk theorem' of repeated game theory in economics (cf. Bowles and Gintis 2011). For a
useful discussion of social behaviour that evolves via reciprocation of benefits, see Sachs et al.
2004.
Despite the attention paid to reciprocal altruism by theoreticians, clear-cut empirical examples in
non-human animals are relatively few (Hammerstein 2003, Sachs et al. 2004, Taborsky 2013). This
is probably because the pre-conditions for reciprocal altruism to evolve- multiple encounters and
individual recognitionare not especially common. However, one possible example is provided by
blood-sharing in vampire bats (Wilkinson 1984, 1990, Carter & Wilkinson 2013). It is quite
common for a vampire bat to fail to feed on a given night. This is potentially fatal, for bats die if
they go without food for more than a couple of days. On any given night, bats donate blood (by
regurgitation) to other members of their group who have failed to feed, thus saving them from
starvation. Since vampire bats live in small groups and associate with each other over long periods
of time, the preconditions for reciprocal altruism are likely to be met. Wilkinson and his colleagues'
studies showed that bats tended to share food with their close associates, and were more likely to
share with others that had recently shared with them. These findings appear to accord with
reciprocal altruism theory.
Trivers (1985) describes an apparent case of reciprocal altruism between non con-specifics. On
tropical coral reefs, various species of small fish act as cleaners for large fish, removing parasites
from their mouths and gills. The interaction is mutually beneficialthe large fish gets cleaned and
the cleaner gets fed. However, Trivers notes that the large fish sometimes appear to behave
altruistically towards the cleaners. If a large fish is attacked by a predator while it has a cleaner in
its mouth, then it waits for the cleaner to leave before fleeing the predator, rather than swallowing
the cleaner and fleeing immediately. Trivers explains the larger fish's behaviour in terms of
reciprocal altruism. Since the large fish often returns to the same cleaner many times over, it pays to
look after the cleaner's welfare, i.e., not to swallow it, even if this increases the chance of being
wounded by a predator. So the larger fish allows the cleaner to escape, because there is an
expectation of return benefitgetting cleaned again in the future. As in the case of the vampire
bats, it is because the large fish and the cleaner interact more than once that the behaviour can
evolve.

5. But is it Real Altruism?

The evolutionary theories described above, in particular kin selection, go a long way towards
reconciling the existence of altruism in nature with Darwinian principles. However, some people
have felt these theories in a way devalue altruism, and that the behaviours they explain are not
really altruistic. The grounds for this view are easy to see. Ordinarily we think of altruistic actions
as disinterested, done with the interests of the recipient, rather than our own interests, in mind. But
kin selection theory explains altruistic behaviour as a clever strategy devised by selfish genes as a
way of increasing their representation in the gene-pool, at the expense of other genes. Surely this
means that the behaviours in question are only apparently altruistic, for they are ultimately the
result of genic self-interest? Reciprocal altruism theory also seems to take the altruism out of
altruism. Behaving nicely to someone in order to procure return benefits from them in the future
seems in a way the antithesis of real altruismit is just delayed self-interest.
This is a tempting line of argument. Indeed Trivers (1971) and, arguably, Dawkins (1976) were
themselves tempted by it. But it should not convince. The key point to remember is that biological
altruism cannot be equated with altruism in the everyday vernacular sense. Biological altruism is
defined in terms of fitness consequences, not motivating intentions. If by real altruism we mean
altruism done with the conscious intention to help, then the vast majority of living creatures are not
capable of real altruism nor therefore of real selfishness either. Ants and termites, for example,
presumably do not have conscious intentions, hence their behaviour cannot be done with the
intention of promoting their own self-interest, nor the interests of others. Thus the assertion that the
evolutionary theories reviewed above show that the altruism in nature is only apparent makes little
sense. The contrast between real altruism and merely apparent altruism simply does not apply to
most animal species.
To some extent, the idea that kin-directed altruism is not real altruism has been fostered by the use
of the selfish gene terminology of Dawkins (1976). As we have seen, the gene's-eye perspective is
heuristically useful for understanding the evolution of altruistic behaviours, especially those that
evolve by kin selection. But talking about selfish genes trying to increase their representation in
the gene-pool is of course just a metaphor (as Dawkins fully admits); there is no literal sense in
which genes try to do anything. Any evolutionary explanation of how a phenotypic trait evolves
must ultimately show that the trait leads to an increase in frequency of the genes that code for it
(presuming the trait is transmitted genetically.) Therefore, a selfish gene story can by definition be
told about any trait, including a behavioural trait, that evolves by Darwinian natural selection. To
say that kin selection interprets altruistic behaviour as a strategy designed by selfish genes to aid
their propagation is not wrong; but it is just another way of saying that a Darwinian explanation for
the evolution of altruism has been found. As Sober and Wilson (1998) note, if one insists on saying
that behaviours which evolve by kin selection / donor-recipient correlation are really selfish, one
ends up reserving the word altruistic for behaviours which cannot evolve by natural selection at
all.
Do theories of the evolution of biological altruism apply to humans? This is part of the broader
question of whether ideas about the evolution of animal behaviour can be extrapolated to humans, a
question that fuelled the sociobiology controversy of the 1980s and is still actively debated today
(cf. Boyd and Richerson 2006, Bowles and Gintis 2011, Sterelny 2012). All biologists accept that
Homo sapiens is an evolved species, and thus that general evolutionary principles apply to it.
However, human behaviour is obviously influenced by culture to a far greater extent than that of
other animals, and is often the product of conscious beliefs and desires (though this does not
necessarily mean that genetics has no influence.) Nonetheless, at least some human behaviour does
seem to fit the predictions of the evolutionary theories reviewed above. In general, humans behave
more altruistically (in the biological sense) towards their close kin than towards non-relatives, e.g.
by helping relatives raise their children, just as kin selection theory would predict. It is also true that
we tend to help those who have helped us out in the past, just as reciprocal altruism theory would
predict. On the other hand, humans are unique in that we co-operate extensively with our non-kin;
and more generally, numerous human behaviours seem anomalous from the point of view of
biological fitness. Think for example of adoption. Parents who adopt children instead of having
their own reduce their biological fitness, obviously, so adoption is an altruistic behaviour. But it is
does not benefit kinfor parents are generally unrelated to the infants they adoptand nor do the
parents stand to gain much in the form of reciprocal benefits. So although evolutionary
considerations can help us understand some human behaviours, they must be applied judiciously.
Where human behaviour is concerned, the distinction between biological altruism, defined in terms
of fitness consequences, and real altruism, defined in terms of the agent's conscious intentions to
help others, does make sense. (Sometimes the label psychological altruism is used instead of real
altruism.) What is the relationship between these two concepts? They appear to be independent in
both directions, as Elliott Sober (1994) has argued; see also Vromen (2012) and Clavien and
Chapuisat (2013). An action performed with the conscious intention of helping another human
being may not affect their biological fitness at all, so would not count as altruistic in the biological
sense. Conversely, an action undertaken for purely self-interested reasons, i.e., without the
conscious intention of helping another, may boost their biological fitness tremendously.
Sober argues that, even if we accept an evolutionary approach to human behaviour, there is no
particular reason to think that evolution would have made humans into egoists rather than
psychological altruists (see also Schulz 2011). On the contrary, it is quite possible that natural
selection would have favoured humans who genuinely do care about helping others, i.e., who are
capable of real or psychological altruism. Suppose there is an evolutionary advantage associated
with taking good care of one's childrena quite plausible idea. Then, parents who really do care
about their childrens' welfare, i.e., who are real altruists, will have a higher inclusive fitness, hence
spread more of their genes, than parents who only pretend to care, or who do not care. Therefore,
evolution may well lead real or psychological altruism to evolve. Contrary to what is often
thought, an evolutionary approach to human behaviour does not imply that humans are likely to be
motivated by self-interest alone. One strategy by which selfish genes may increase their future
representation is by causing humans to be non-selfish, in the psychological sense.

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Related Entries
game theory: evolutionary | natural selection | natural selection: units and levels of | sociobiology
Copyright 2013 by
Samir Okasha <samir.okasha@bristol.ac.uk>