Altruism
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.
Why do most species live in groups?
There are many reasons for species tending to occur in groups:
• increased chance of surviving:
probability of being predated lower on average, most
predation occurs on animals at the periphery of the group
• increased chance of finding a mate:
higher density in groups than if scattered
• increased chance of finding food and feeding -
some keep watch while others feed, jobs exchanged over time,
everyone gets more time to feed. There are costs, as well as
benefits...
What is Altruism?
• Rooted in the Latin word alter – meaning other
• Altruism – means “living for others”
• Key component – selflessness – an unselfish regard
for the welfare of others
• Altruism : “refers to acts that are carried out
voluntarily by individuals who have no concern for
themselves and who have no expectation of any kind
of reward
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 ) standard)
Direct fitness: reproductive success based on
your own reproduction.
Indirect fitness: reproductive success based
on the helping of related individuals (above
what they otherwise would have raised).
Inclusive fitness: total fitness due to both
direct and indirect fitness.
• Inclusive fitness = direct fitness + indirect fitness
You are increasing your own genetic fitness
by helping, even if your own direct fitness
suffers.
Natural selection favors these genes, and is
known as indirect or kin selection.
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.
• In many species of social insects, workers forgot
reproduction entirely (they are sterile) in order to help
raise their sisters (Wilson, 1971).
Vampire bats will regurgitate and feed blood that
they have collected from their prey to a hungry
conspecific (A member of the same species)
(Wilkinson, 1990).
Beldings 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).
So why call?
Alarm calling
Meerkat sentinels (guard )
responsible for alerting others who may be busy foraging for food
to the presence of predators
Best examples come from eusocial insects
Bees, termites, ants
Suicide in bees, etc.
Non-breeding workers
Theories of Altruism.
• Kin Selection
• Proposes that we are more likely to act altruistically when it comes to saving
our relatives
• Reciprocal Altruism
• Suggests that helping in the short run increases the probability that our
genes will be protected in the future.
• What goes around, comes around
• Helps explain non-kin helping behavior
Group Selection
• This perspective argues that groups consisting of cooperative members are
more likely to survive and pass on their genes than groups composed of
selfish members
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 one’s
‘Inclusive Fitness’.
Evolutionary fitness is measured by the numbers of
copies of ‘your’ genes in the subsequent generation
compared to those of others.
Proportion shared
To assess the contribution
Self 100%
to inclusive fitness made by father 50%
mother 50%
a relative, you need to full sibling 50%
half sibling 25%
know the coefficient of offspring 50%
relationship grandparents 25%
niece/nephew 25%
(the fraction of your uncle/aunt 25%
first cousin 12.5%
genome shared from
ancestors in common)
Why should some individuals
help others at the expense of
their own 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.
‘Hamilton’s Rule’ = r B > c
• where r=coefficient of relatedness, B = benefit to the recipient, c = cost to the
giver.
• Hamilton’s rule states that a trait is favored by natural selection
if the benefit to others, B, multiplied by relatedness, R, exceeds
the cost to self, C
Altruistic Selfish
behaviour behaviour
can evolve in occurs
this region only in
this
region
monkey: give an alarm call and will earn a cost (being
eaten) at the benefit of the others
Monkey remain silent, where some of the other
members will be eaten while the actor escapes
unharmed
From the previous examples
If the beneficiaries of this act of altruism are close
relatives, it would actually benefit the inclusive
fitness of that individual to give the call and save
their kin rather than themselves.
Because relatives will have a higher tendency to
share similar genetic material
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).
Warning colouration in insects, e.g. the
larvae of the monarch butterfly. It contains
toxins poisonous to bird predators.
The first brightly coloured larvae would
have attracted predators, and suffered total
loss of individual fitness. Siblings are
‘protected’ by bird learning. Inclusive fitness
increases through kin selection.
2. Reciprocal Altruism
• Proposed by Trivers (1971).
• Natural Selection may create psychological mechanisms designed to
deliver benefits even to non-relatives, provided that such actions lead to
reciprocal beneficial actions in the future.
• This is not necessarily limited to the same species e.g. cleaner fish.
• If the benefit received is larger than the cost incurred (gained), 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).
Reciprocal altruism
A will help B, if B will help A in the future
Reciprocal altruism can evolve only under a restricted set of
conditions
Need:
-repeated interactions
-individual recognition
Kin Recognition.
• It is important to be able to recognise kin, as the costs involved in mistaking
another individual's offspring for one’s 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.
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.
• Kin selection does not explain observed incidences of animals helping non-
relatives for example:
• Unrelated chimpanzees come to one another’s 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.
• Rescuing unrelated individuals (and even animals).
• Sacrificing their lives for moral or ethical principles.
• How can such behaviours be explained?
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 non-repeated interactions when
gains are small.
• Strong reciprocity is the tendency 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.
Group selection
1962, Wynne-Edwards.
Group selection is controversial because conditions that lead
to cooperation among unrelated individuals are very
restrictive in evolutionary terms.
Self-interest is the dominant force.
Ants who are workers do not reproduce but without their
sacrifice the ants would not be successful.
Lemmings migrate away when food is scarce, risking their
lives, so resources would be available for others of the
species.
There are two recent approaches to group selection
that could work:
1)a population exists as a set of groups. Among those
groups, isolated, selfish subgroups must go extinct
faster than selfishness
2)Populations come together and separate into
subgroups. Some subgroups have different rates of
survival and/or reproduction.
Why doesn't group selection work?
Prone to invasion by
Selfish gene.
