Evolutionary Psychology,

Lecture 3.
Altruism and Co-operation

Learning Outcomes.
At the end of this session you should be
able to:
1. Explain what is meant by the terms kin
selection and reciprocal altruism.
2. Discuss kin selection and reciprocation
explanations
for
human
and
animal
cooperative / altruistic behaviours.

Thoughts For the Day.

Let us try to teach generosity and altruism because
we are born selfish Richard Dawkins (1976).
ethics, morality, human conduct, and the human
psyche are to be understood only if societies are
seen as collections of individuals seeking their own
self interest. R.D. Alexander (1987).
Scratch an altruist and watch a hypocrite bleed.
Ghiselin (1974).

Altruism.
Altruism refers to an individual acting in a way that
will decrease its own survival chances, but improve
the survival chances of another individual.
The Darwinian perspective emphasising survival of
the fittest gave the impression that selfishness was
the norm.
Pioneering work involving the study of animals living
in social groups in fact revealed that co-operation
and altruism are just as natural as selfishness.
If co-operation and altruism have evolved, then they
must have some adaptive benefits, researchers have
analysed the conditions under which adaptations for
engaging in such behaviour can be expected to
evolve.

Examples of Animal
Altruism.
Vampire bats will regurgitate
and feed blood that they have
collected from their prey to a
hungry
conspecific
(Wilkinson, 1990).
Ground squirrels will warn
others of the presence of a
predator, even though making
such a call may draw the
attention of the predator to
itself (Sherman, 1977).
In many species of social
insects,
workers
forgo
reproduction entirely (they
are sterile) in order to help
raise their sisters (Wilson,
1971).

Theories of Altruism.
1. Kin Selection (Proposed by Hamilton, 1964).
By helping relatives to reproduce (even at the cost
to your own reproductive success) then your shared
genes can spread. Assisting a close relative thereby
increases ones Inclusive Fitness.
Using mathematical modelling, Hamilton showed
that an altruistic gene can spread through the
population if it causes an individual to help a
relative, whenever the cost to the individual is offset
by the reproductive benefit gained by the receiver.
Hamiltons Rule = r B>c
where r=coefficient of relatedness, B = benefit to
the recipient, c = cost to the giver.

Kin Selection in Action.

Ground squirrels do not give an alarm call every
time a predator approaches. They only do so when
there is a large proportion of their relatives within
earshot (Sherman, 1977).
Vampire bats are much more likely to share their
food
with
relatives
than
with
non-relatives
(Wilkinson, 1990).
This theory explained the most puzzling phenomena
- that of the sterile insects - by a genetic quirk they
are more related to their sisters than to their
mothers or daughters (Trivers & Hare, 1976).

Kin Recognition.
It is important to be able to recognise kin, as the costs
involved in mistaking another individuals offspring for
ones own are high, and the benefits few.
Offspring recognition should evolve more often in
colonial species, as there is a high risk of misdirecting
parental care.
Examples.
Bank swallows (colonial) do not accept strange chicks
whereas rough-winged swallows (solitary) do.
Herring gulls (colonial ground-nesting) recognise
offspring and refuse strange chicks, but Kittiwakes
(colonial cliff-nesting) do not recognise offspring and
accept substitute offspring.

Kin Recognition in
Gulls.

Data from Alcock, 1993

Kin Selection in
Humans.
Studies
amongst
diverse
human
populations
consistently support the existence of kin selection, some
examples (cited in Barrett et al., 2002) are as follows:
Food sharing is more common amongst close relatives.
Political alliances between kin are more stable than
those formed between distantly related, or unrelated
individuals and involve less preconditions.
The passing on of wealth to lineal descendants
(excluding spouses) is far more common than giving to
less closely related or unrelated individuals.
Close relatives are preferentially sought out in times of
need and such help is less likely to be reciprocal.
Relatives typically receive more expensive presents.

How Much Pain Will You

Suffer
For
Your
Kin?
In an interesting experiment Fieldman et al., (cited

in Barrett et al., 2002) asked participants to

maintain a painful position. The longer they held the
position the more money they would earn.
In different conditions participants could earn
money for individuals differing in relatedness:
Themselves.
Parent or sibling.
Grandparent / niece / nephew, a cousin.
Unrelated friend.
The duration of maintaining the painful position
varied as a direct proportion of relatedness, with
more pain being sustained for closer relatives.

Facial Similarity and Trust?

DeBruine (2002) argued that animals should be
sensitive to cues of genetic relatedness when making
altruistic decisions.
In humans such decisions may be based around facial
appearance.
Participants played a computerised game of trust in
which they had to decide whether or not to share
money with an individual.
They were shown faces of their 'opponents' which were
either facially different to themselves, or whose faces
had been morphed to resemble their own.
Participants showed significantly
behaviour when playing against
resembled themselves.

more 'trusting'
opponents that

Human Adoption.
The adoption of unrelated children has been cited as
evidence against kin selection as helping to rear unrelated
children will not produce genetic benefits to the giver.
However, Silk (1990) observed that among Polynesian
cultures, a substantial number of adopters cared for
children who were cousin equivalents or closer. Families
who had adopted children that were unrelated tended to
be agricultural families needing extra help.
Similarly, in Chicago Stack (1974) reported that the
majority of foster children were adopted by kin.
Adopting unrelated children is a recent Western
phenomenon. Alcock (1993) argues that the urge to
produce children and look after them is so beneficial in
reproductive terms that it has become deeply ingrained.

Problems for Kin

Selection.
Kin selection does not explain observed incidences of
animals helping non-relatives for example:
Unrelated chimpanzees come to one anothers aid
when threatened (de Waal & Luttrell, 1988).
Vampire bats will feed non-relatives (Wilkinson, 1990).
Humans often engage in apparently altruistic acts such
as:
Giving blood.
Donating to charity.
Forgoing reproduction.
Rescuing unrelated individuals (and even animals).
Sacrificing their lives for moral or ethical principles.
How can such behaviours be explained?

2. Reciprocal
Altruism.
Proposed by Trivers (1971).
Natural
Selection
may
create
psychological
mechanisms designed to deliver benefits even to nonrelatives, provided that such actions lead to
reciprocal beneficial actions in the future.
you scratch my back.
This is not necessarily limited to the same species
e.g. cleaner fish.
If the benefit received is larger than the cost
incurred, then individuals who engage in such
behaviour will out-reproduce those who do not.
Eg, in vampire bats, an individual will share food with
a conspecific (whether related or not) if the other has
shared food with that individual in the past
(Wilkinson, 1990).

Conditions Under Which

Reciprocation Flourishes.
1. Individuals must associate for long-enough
periods of time to develop reciprocal interactions.
2. The likelihood of one individual performing some
social exchange with another should be predicted on
the basis of their past associations.
3. The roles of giver and receiver should reverse at
least once.
4. The short-term benefits to the recipient are
greater than the costs to the donor.
5. Givers should be able to recognise and expel
cheaters from the system.
Do such conditions apply to human social
interactions?

Modelling Human
Social Exchanges.
Game Theory was developed by the mathematician
von Neumann and the economist Morgenstern in the
1940s in an attempt to model the behaviour of
individuals in economic and adversarial situations.
Maynard-Smith (1982) adapted it to model cooperation and competition in the social world.
Within game theory, individuals behave rationally
and choose the action that yields the highest payoff.
Human social interactions can be mapped into game
settings, as one individual can benefit from the
actions of others at little cost to themselves.
There are several forms:

Unscrupulous Diner
Scenario.
A group of diners agree to divide the restaurant bill
equally, most co-operate by choosing similar priced
meals but an individual can take advantage by ordering
the most expensive meal, as the cost will be absorbed
by the whole group (Glance & Huberman, 1994).
In a one-off situation in a large social group it pays to
cheat, however in a small group who meet regularly,
such defection will be noticed and punished.
Reciprocal social exchange has mutual costs/benefits
but one person can always benefit more than another if
they cheat - i.e. receive an act but do not reciprocate.
This constitutes a formidable barrier to the evolution of
social exchange.

Prisoners Dilemma.
Described by Axelrod & Hamilton (1981). It is a game in
which mutual co-operation benefits both players, but a
cheat can gain a higher pay-off.
It is often described as a hypothetical situation in which
two individuals have committed a crime, and are being
held for questioning in separate cells, they are unable
to communicate.
It is in the best interests of both to say nothing, as the
evidence is such that both may only receive a light
sentence.
However, they are being questioned separately, and the
lawyer offers both freedom if they implicate the other
in the crime.
When the game is played once or only a few times then
the best strategy is to inform on the other (defect).

Prisoners Dilemma Payoff

Matrix
Player B
Co-operates

Player B
Defects

Player A
Co-operates

R=3 year sentence

each.
Reward for mutual
cooperation

S= long sentence
for A, freedom for
B.
Sucker's payoff

Player A
defects

T= freedom for A,
long sentence for B.
Temptation to
defect

P=10 year sentence

each.
Punishment for
mutual defection

Which Strategy is
Best?
Real life social exchanges may occur more than once and
individuals will probably remember those who have
defected (cheated) in the past.
Axelrod (1984) organised a tournament in which various
computer programs played repeated prisoners dilemma
games. A program called 'tit for tat' was a clear winner.
This simple strategy always cooperated on the first
move, and then copied its opponents move on every
subsequent move.
This is an Evolutionary Stable Strategy (ESS) - i.e. once
established it cannot be displaced by another strategy.
In real life it pays to cheat in a one-off exchange
encounter but if there is a likelihood that you will
encounter the same person more than once, then mutual
cooperation will serve both parties the best.

Indirect
Reciprocation.
Trivers (1971) argued that an altruistic act need not
necessarily be reciprocated by the person directly
assisted but can be returned indirectly from other
individuals.
E.g if you advertise yourself as an altruist then
individuals will be more favourably inclined to deal with
you in future social exchange situations.
This may explain blood donation, giving to beggars, and
donating to charities.
It had been claimed that such actions indicate that
human behaviour is immune form evolutionary analysis
and demonstrates a pure form of altruism.
Alexander (1987) suggested that giving blood is a very
good way of demonstrating your altruism at only a
modest cost.

Evidence for Indirect

Reciprocation.
We would maybe predict that individuals will not donate
blood or give to charity unless their actions are made
known i.e. by wearing a sticker or badge indicating their
actions.
Low & Heinen (cited in Alcock, 1993) reported that
students are significantly more likely to give to charity if
they receive a pin or tag that advertises their
participation.
Mulcahy (1999) observed who gave money to beggars and
then interviewed mixed-sex couples after the male had
donated money.
Males at the early stage of a relationship were more likely
to give than those in a long-established relationship.
Thus the act of giving when accompanied by a female is
not simply to do with impressing a current partner, but
demonstrating one's generosity to a potential partner.

An Alternative
Theory.
Gintis et al., (2003) argues that kin selection and
reciprocity theories do not explain why cooperation is
frequent amongst unrelated individuals in nonrepeated interactions when gains are small.
Strong reciprocity is the predisposition to cooperate
with others, and to punish those who violate the
norms of cooperation, at some personal cost, even
when such costs may not be repaid.
In support, Fehr & Gchter (2002) showed that when
asked to play a game for monetary reward under 'no
punishment' or 'punishment conditions, punishment
of non-cooperators substantially increased the
amount that groups invested for the good of the
group.

Additional Evidence For

Strong Reciprocity.
The concept of 'fairness' lies at the heart of many
human social interactions and can be modelled using
the 'Ultimatum game'.
Here a participant is given a sum of money and told
they can keep it provided that they split the sum
with another individual.
The participant has to make a one-off offer between
0 -100% of the total sum to the other person.
If the second person agrees to the offered sum then
both keep these amounts; if they reject the offer
then both receive nothing. No haggling is allowed.
According to one-off game theory exchanges we
would expect that the first participant would offer a
sum of well below 50% and that the receiver should
accept any sum as anything is better than nothing.

The Ultimatum Game in

Action.
However when this game is played it is typically found
that individuals offer around 50%, and more than half of
receivers do not accept offers less than 20% (Sigmund
et al., 2002).
Individuals do not behave completely selfishly but place
a high value on fair outcomes. Receivers are prepared to
accept smaller 'gifts' under the following conditions:
The giver is chosen by better performance on a quiz.
The givers offer is randomly selected by a computer.
Several responders compete to accept a proposers offer.
Sigmund et al., (2002) proposed that our emotional
apparatus has been shaped over millions of years of
small group living in which it is hard to cheat more than
once and where we expect conspecifics to notice our
actions and remember them.