Evolution and The Social Contract: The Tanner Lectures On Human Values
Evolution and The Social Contract: The Tanner Lectures On Human Values
Brian Skyr ms
Delivered at
[49]
Prisoner’s Delight
Cooperate Defect
Cooperate 3 1
Defect 2 0
The entries in the table represent the payoffs of Row’s strategy when
played against Column’s strategy. If your partner cooperates, you are bet-
ter off if you do as well, for a payoff of 3 rather than 2. If your partner
defects, you are still better off cooperating for 1 rather than 0, although
your partner does even better with 2. So no matter what your partner does,
you are better off cooperating. Your partner is in the same situation, and
reasons likewise. It is easy to cooperate.
We can change the story a little bit to fit a different kind of situation.
Perhaps if your partner defects, your attempt at cooperation is counter-
productive. You are better off defecting. This change gives us the game
known as the Stag Hunt:
Stag Hunt
Cooperate Defect
Cooperate 3 1
Defect 2 2
In the Stag Hunt, what is best for you depends on what your partner
does. If you both cooperate, you are both doing the best thing given your
partner’s action—likewise if you both defect. Cooperation is more dif-
ficult. It is an equilibrium, but not the only one.
Another modification of the situation calls for a different game. Sup-
pose that defecting against a cooperator actually pays off. Then we have
the Prisoner’s Dilemma:
Prisoner’s Dilemma
Cooperate Defect
Cooperate 3 1
Defect 4 2
Your optimal act again no longer depends on what your partner does.
Now it is to defect. Cooperation is hard.
Each of these games is a reasonable model of some social interactions.
Two men sit in a rowboat, one behind the other. Each has a set of oars.
They have been out fishing, and a hot dinner awaits them across the lake. If
one doesn’t row for some reason, the other will row to get them there; if one
does row, the other prefers to row also to get home faster. This is Prisoner’s
Delight. Cooperation is easy. Now change the picture. They sit side by side,
and each has one oar. One man rowing alone just makes the boat go in
circles. This is a Stag Hunt.1 Back to the first rowboat with two sets of oars,
but take away the hot dinner on the opposite shore and suppose that the
men are very tired. They could camp on this shore for the night, although
they would prefer to get back to the opposite shore. But either prefers not
to row no matter what the other does. This is a Prisoner’s Dilemma.
Consider any reasonable adaptive dynamics operating on a large popu-
lation of individuals paired at random to play one of these games. Then in
Prisoner’s Delight cooperation takes over the population, in Prisoner’s Di-
lemma defection goes to fixation, and in Stag Hunt one or the other may
prevail depending on the initial composition of the population.
There is an enormous literature devoted to explaining the evolution
of cooperation in the Prisoner’s Dilemma, while the other two games are
relatively neglected. Everyone wants to crack the hardest problem—the
evolution of altruism.2
All these accounts of cooperation in Prisoner’s Dilemma either (1) use
an interaction that is not really a Prisoner’s Dilemma or (2) use pairings to
play the game that are not random. It must be so.3 Suppose the interaction
is a Prisoner’s Dilemma, and pairings are random in a large population.
Then cooperators and defectors each have the same proportion of part-
ners of each type. Defectors must on average do better than cooperators.
Replicator dynamics increases the proportion of the population of the
type that does better, and that’s all there is to it.
But if nature somehow arranges for positive correlation—for coopera-
tors to meet cooperators and defectors to meet defectors more often than
they would with random matching—then it is possible for cooperators to
do better than defectors. The point is obvious if we consider perfect correla-
tion. Then the relevant comparison in payoffs is not vertical but diagonal:
1. This is David Hume’s famous example from the Treatise of Human Nature. For more
game theory in Hume, see Vanderschraaf 1998.
2. Binmore (1994) devotes a chapter titled “Squaring the Circle in the Social Sciences” to
attempts to justify cooperation in the one-shot Prisoner’s Dilemma.
3. This is the “Iron Rule of Selfishness” of Bergstrom (2002). See also Eshel and Cavalli-
Sforza 1982.
Prisoner’s Dilemma
Cooperate Defect
Cooperate 3 1
Defect 4 2
4. The use of discounted repeated games to explain cooperation in Prisoner’s Dilemma is
already to be found in Luce and Raiffa 1957, with no claim of originality. John Nash is reported
to have invented the explanation in conversation.
repeated games. Suppose that the probability that the Prisoner’s Dilemma
will be repeated another time is constant. This (somewhat far-fetched)
idealization of geometrical discounting of the future allows us to sum an
infinite series and compute the expected payoffs of strategies in the large
repeated game. For simplicity, we consider only two strategies in the re-
peated game, Always Defect and Tit for Tat. A Tit for Tat player initially
cooperates and then does what was done to him in the preceding round,
and Always Defect is self-explanatory. Two players remain matched for
the whole repeated game, but this restrictive assumption can be relaxed
in more complicated “community enforcement” models (Sugden 1986;
Kandori 1992; Milgrom, North, and Weingast 1990; Nowak and Sigmund
1998).
Since the payoffs on each individual play are those of Prisoner’s Di-
lemma, the strategies in the repeated games must induce a correlation be-
tween individual plays of Cooperate and Defect if cooperation is not to
be driven to extinction. The presence of Tit for Tat players in the popula-
tion is this correlation device. They always cooperate with each other and
quickly learn to defect against defectors.
What is the larger game in which plays of Prisoner’s Dilemma are em-
bedded? Using the version of Prisoner’s Dilemma given before and prob-
ability of another trial as six-tenths, we get:
This is a Stag Hunt. There are two stable equilibria, one where everyone
always plays Tit for Tat and one where everyone always plays Always De-
fect. Which one you get depends on initial population proportions. In our
example an initial population split equally evolves to universal defection.
Sober and Wilson (1998) direct our attention to group selection. Con-
sider the haystack model of Maynard Smith (1964). In the fall, farmers cut
hay and field mice randomly colonize the haystacks. In the haystacks they
play the Prisoner’s Dilemma and reproduce according to the payoffs. In
the spring the haystacks are torn down, the mice scatter, and the cycle is
continued. If there are enough generations of mice in the life of a haystack,
then it is possible for cooperators to do better on average than defectors.
This is because differential reproduction within haystacks creates positive
correlation in the population. In haystacks colonized by cooperators and
5. There are pioneering papers by Pollock (1989), Nowak and May (1992), and Hegsel-
mann (1996). These are models where the interaction is Prisoner’s Dilemma, or in the case of
Nowak and May at a bifurcation between Prisoner’s Dilemma and Hawk-Dove. Ellison (1993,
2002) discusses local interaction models of the Stag Hunt in which the <Defect, Defect> equi-
librium is risk-dominant. In his models it is the noncooperators who can invade and quickly
take over. The differences between Ellison’s models and ones in which cooperators can invade
and take over are discussed in Skyrms 2004.
Prisoner’s Dilemma
Cooperate Defect
Cooperate 3 1
Defect 4 2
Stag Hunt
Cooperate Defect
Cooperate 3 1
Defect 2 2
6. For other models of correlation induced by partner choice, see Wright 1921, where the
literature begins; Hamilton 1971; Feldman and Thomas 1987; Kitcher 1993; Oechssler 1997;
Dieckmann 1999; and Ely 2002.
7. For an experiment in a public goods–provision game with voluntary association, see
Page, Putterman, and Unel 2005.
place in a petri dish. Theoretical analysis by Durrett and Levin (1994) and
Iwasa, Nakamaru, and Levin (1998) is a complement to the local interac-
tion model of evolution of cooperation. If we put this case in the frame-
work of Eshel, Shaked, and Sansone of the last section, we find that a large
interaction neighborhood and a small imitation neighborhood robustly
favor the evolution of spite (see Skyrms 2004).
Group-selection models are not without their spiteful aspects. Sup-
pose that the farmer never tears down the haystacks—the population is-
lands that they represent remain isolated. Then for a mouse, its haystack
becomes its world, and this makes all the difference (see Gardner and
West 2004). Its haystack’s population becomes a population unto itself,
within which evolution takes place. The carrying capacity within a hay-
stack is limited, so we are dealing with a small, finite population. This,
in itself, induces negative correlation even if pairs of individuals form at
random within the haystack, because an individual does not interact with
herself. This effect is negligible in a large population but can be significant
in a small population. (For a transparent example, consider a population
consisting of four individuals, two C and two D. Population frequencies
are 50-50, but each type has probability 2/3 of meeting the other type and
1/3 of meeting its own.)
If a defector is introduced into a haystack full of cooperators—a mu-
tant or a migrant—he can cause problems. If the interaction is Prisoner’s
Dilemma, defectors will, of course, take over. But in small populations,
with some versions of Stag Hunt and even Prisoner’s Delight, defectors
can still invade as a result of negative correlation.
For any positive value of e, the following is a version of Prisoner’s De-
light—an individual prefers to cooperate no matter what the other does:
8. For example, suppose 1 defector is introduced into a population of N cooperators. In-
dividuals pair at random. Since the defector cannot interact with himself, he always pairs with
a cooperator for a payoff of 2. Cooperators pair with the defector with probability (1/N) and
with other cooperators with probability (N-1)/N, for an average payoff of [(N-1)/N]*2 + e. So
if e < (2/N), a spiteful mutant does better than the native cooperators.
Bargaining
Prisoner’s Delight, Stag Hunt, and Prisoner’s Dilemma are not the only
games that raise issues central to the social contract. We could separate
the issues of cooperating to produce a public good and deciding how that
good is to be divided. This is the philosopher’s problem of distributive
justice, and it brings bargaining games to center stage (Braithwaite 1955;
Rawls 1957; Sugden 1986; Binmore 1994, 1998, 2005; Gauthier 1985, 1986).
Consider the simplest Nash Bargaining game. Two players have bot-
tom-line demands for a share of a common good. If the demands exceed
the available good, agreement is impossible and players get nothing. Oth-
erwise, they get what they demand. We simplify radically by assuming
that there are only three possible demands: one-third, one-half, and two-
thirds. Evolutionary dynamics in a large random-encounter environment,
with and without persistent random shocks, is well studied.
Allowing differential reproduction to carry a population to equilib-
rium, there are two possibilities. The population may reach an egalitar-
ian consensus, where all demand one-half. Or it may reach a polymorphic
state, where half the population demands two-thirds and half the popu-
lation demands one-third (Sugden 1986). Greedy players get their two-
thirds half the time; modest players get their one-third all the time. This
inefficient polymorphism wastes resources, but it is evolutionarily stable
and has a significant basin of attraction. Persistent shocks can allow a
population to escape from this polymorphic trap and favor the egalitarian
norm in the very long run, but the inefficient polymorphism remains a
possibility for medium-run behavior.9
However, the effect of correlation mechanisms is rarely discussed in
connection with Nash Bargaining. If correlation plays an important role
in producing a surplus to be divided, might it not also play an important
role in deciding how the division takes place? Positive correlation of de-
mand types obviously favors the egalitarian solution. Those who ask for
9. Most of what we know about this is due to Peyton Young. See Young 1993a, 1993b,
1998; and Binmore, Samuelson, and Young 2003.
equal shares do best when they meet each other. Negative correlation is
more complicated. Greedy players who demand two-thirds do very well
if paired with modest players who ask for one-third, but not well if paired
with those who ask for half. If negative correlation initially allows greedy
players to outreproduce all others, it cannot be maintained because they
run out of modest players. But a greedy-modest polymorphism is a real pos-
sibility if the negative correlation is of the kind that sufficiently disadvan-
tages the egalitarians. Possibilities are multiplied if we allow more demand
types. It should be of interest to look at specific correlation mechanisms.
If we allow individuals to bargain with neighbors on a spatial grid,
islands of egalitarianism spontaneously form. This generates positive cor-
relation, and if individuals emulate their most prosperous neighbors, egal-
itarianism takes over the population. This is a very rapid process, as other
types who interact along the edges of the egalitarian islands are quickly
converted to asking for half ( J. Alexander and Skyrms 1999; J. Alexander
2007; Skyrms 2004). Equal sharing is contagious.
If we add costless signaling to a large-population random-encounter
evolutionary model of bargaining, complicated correlations arise and then
fade away. Cooperators establish a positive correlation with cooperators.
Greedy types establish a negative correlation with themselves. Although
these correlations are transient, their effect is that the basin of attraction of
the egalitarian equilibrium is greatly enlarged (see Skyrms 2004).
Axtell, Epstein, and Young (2006) investigate a related model where
individuals have one of two “tags” and can condition their action on the
tag of their partner in a bargaining game. But there is a different dynamics.
Instead of evolution by replication or imitation, they consider a rational-
choice model. Things may fall out in various ways—here is one. When
interacting with those having the same tag, individuals share alike. But
in interactions between tags, one tag type becomes greedy and always de-
mands two-thirds and the other becomes modest and always demands
one-third. In this equilibrium tags are used to set up both positive and
negative correlations between behaviors. Both correlations are perfect: de-
mand-half behaviors always meet themselves, and the other two demand
behaviors always meet each other. The result is egalitarianism within tag
types and inegalitarian distribution between types. Axtell, Epstein, and
Young see this as a spontaneous emergence of social classes.
We can also see the spontaneous emergence of social classes in a dy-
namic social network (Skyrms 2004). The classes are stabilized by rational
choice but destabilized by imitation. Depending on the details and timing
of the dynamics, the social network may end up egalitarian or with a class
structure.
Division of Labor
So far, negative correlation has played a rather sinister role in this story.
It is not always so. In cooperating to produce a common good, organisms
sometimes discover the efficiency of division of labor, and find a way to
implement it. Modern human societies are marvels in their implementa-
tion of division of labor; so are the societies of cells in any multicellular or-
ganism. On the most elementary level, we can suppose that there are two
kinds of specialists that an individual can become, A and B, and that these
specialists are complementary. On the other hand, an individual might
not specialize at all but rather, less efficiently, do what both specialists do.
This gives us a little division-of-labor game:10
Division of Labor
Specialize A Specialize B Go It Alone
Specialize A 0 2 0
Specialize B 2 0 0
Go It Alone 1 1 1
Groups Revisited
Individuals sometimes form groups that have a permanence and unifor-
mity of interaction with other groups that qualified them to be thought of
as individuals. This happens at various levels of evolution. We ourselves are
such groups of cells. And humans participate in various social corpora—
teams, states, ideological groups—that interact with others.
10. For analysis of different division-of-labor games, motivated by evolution of coviruses,
see Wahl 2002.
11. One population signaling models face is when an individual interacts with another
who sends the same signal. See Skyrms 2004.
How such superindividuals are formed and hold together (or don’t)
is a central issue of both biology (R. Alexander 1979, 1987; Buss 1987;
Maynard Smith and Szathmary 1995; Frank 1998, 2003) and social sci-
ence. There is no one answer, but answers may involve both elements of
cooperation and elements of spite. One important factor is the punish-
ment of individuals who act against the interests of the group. A large
experimental literature documents the willingness of many individuals to
pay to punish “free riders” in public goods–provision games, and shows
that such punishment is able to stabilize high levels of cooperation.12 This
is also an important finding of Ostrom’s field studies of the self-organized
government of commons (1990). Costly punishment is, from an evolu-
tionary point of view, a form of spite—although it is not called by that
name in the literature.
The name may strike the reader as overly harsh in the case of the mod-
est, graduated punishments found in Ostrom’s field studies of successful
cooperative collective management. But in tightly organized superindi-
viduals, punishment can be draconian. Totalitarian regimes or ideologies
classify those who violate social norms as traitors or heretics. They may be
stoned to death. They have been burned at the stake. The righteous people
carrying out such acts no doubt believe that they are engaged in “altruistic
punishment.” We need also to think about the dark side of punishment.
When groups that can operate more or less like superindividuals have
been formed, the interactions of the superindividuals themselves are also
liable to the effects of positive and negative correlation described above.
They can cooperate to produce a common good, or not. Their interactions
may exemplify spite—not only in behavior, like the bacteria, but also in
the full psychological sense of the word.
Repeated interactions, alliances, local interaction on a geographical
landscape, signals, tags, and network formation all play a role. Division
of labor is facilitated by trade networks, and trade may promote both the
good and the bad sides of negative correlation.
The negative correlation conducive to spite in small populations
may take on a larger significance when we consider interactions between
groups. A local population of six interacting nations is perhaps more plau-
sible than a local population of six interacting mice.
12. For instance, see Ostrom, Walker, and Gardner 1992; and Fehr and Gachter 2000,
2002. Costly punishment is already implicit in the behavior of receivers in ultimatum-game
experiments from Güth, Schmittberger, and Schwartze (1982) to Henrich et al. (2004).
13. But theorists disagree about the nature of rational choice. Rawls minimizes the maxi-
mum loss; Harsanyi maximizes the expected payoff.
14. The theorist tells you what that choice will be.
15. In large populations, expected fitness can then be calculated using population propor-
tions in place of the subjective probabilities of rational choice theory.
Notes
References