Altruism, Kin Selection, and Cooperative Breeding

Altruism,
Kin Selection,
and Cooperative
Breeding
• Natural selection acts on individuals.
• Traits like behavior do not evolve to benefit the species.
• Traits that are beneficial to individual fitness are favored and spread
through populations and species over time.
If an individual behaved in such a way that

decreased its fitness but helped the species, what
would happen to the genes for that trait over time?
These genes would not be passed on.

This behavior would not evolve.
So, how can helping behavior evolve?

This is an evolutionary puzzle!
We can classify interactions based on their effects
on the fitnesses of interactants:
Effect on Fitness of
Interaction Actor Recipient
Mutualism + +
Self-beneficial
Selfish behavior + -
Spiteful behavior - -
Self-sacrificing
Altruism - +
Mutualism: behavior has positive effect on fitness
of actor and recipient
Bluegill sunfish males care
for eggs
Nests are clumped in groups

to protect against predators
If a male chases off a Bluegill sunfish
predator, this benefits

himself & neighbors We can explain this kind of behavior
by individual selection
Selfish behavior: behavior has positive effect on
fitness of actor at a cost to recipient’s fitness
Bluegill sunfish males care

for eggs
Nests are clumped in groups

to protect against predators
Bluegill sunfish If a male eats the eggs in

Selfish behavior can easily be another male’s nest,
explained by individual selection benefits self; harms other
Altruism: behavior that has a negative effect on the
fitness of the actor, but benefits the recipient
An evolutionary puzzle
Biological definition of altruism: by behaving

altruistically, an individual reduces the number of offspring
it is likely to produce itself, but boosts the number that
other individuals are likely to produce
Belding’s ground squirrel’s “terrestrial predator” alarm call

increases chance of caller being detected by predator
*fitness cost to altruist
Recipients of call benefit by being alerted to predator.

*fitness benefit to recipient
Altruism: behavior has negative effect on fitness of
actor, but benefits recipient
Females with a sister or mother (but no direct offspring)
alarm call as often as females with offspring, and more
than females with no relatives
Is this altruism?
Most biologists agree: giving aid to offspring is parental care and not
altruism, while giving aid to other relatives is altruism
To generate a definition of altruism that includes non-offspring relatives, we need
to make some distinctions between categories of fitness
Recall fitness:
FITNESS = reproductive success of an individual, relative to other members of
the species
DIRECT FITNESS: a measure of an individual’s reproductive (genetic)

success based on the number of its offspring that survive to reproduce
INDIRECT FITNESS: a measure of the genetic success of an altruistic

individual based on the number of the relatives (genetically similar individuals)
that the altruist helps to reproduce.
This brings us to the concept of kin
selection
Kin selection: acting in favor of individuals
that share some of your genes, at a cost to
yourself.
Inclusive fitness = a total measure of an individual's

contribution of genes to the
next generation due to both direct and
indirect selection
Inclusive fitness = direct fitness + indirect fitness

Hamilton’s rule Giving aid to non-descendant kin (altruism)
will evolve if:
Altruistic trait must be
rB > C HERITABLE otherwise how
can it evolve?
r = the genetic relatedness of the recipient to the actor

W. D. Hamilton
B = the additional reproductive benefit gained by the
recipient of the altruistic act
C = the reproductive cost to the individual performing the act

(the altruist)
Coefficients of relatedness (r)
Mom Dad Common coefficients (r)

parent- Comparison r
.5 of .5 of offspring
}
Parent & kid 1/2
genes genes related by
0.5 Sibs 1/2
Identical twins 1
Offspring Sibling Half sibs 1/4
{
Full siblings related by 0.5
Cousins
Aunt to her nephew or

1/8
1/4
niece
Male turkeys often court females in
coalitions (2-4 males)
Display jointly, cooperatively ward
off other males
Dominant males have higher reproductive success

than solo males
Subordinates have 0 reproductive success
Why should subordinates make
this sacrifice?
Beneficial if they are related to dominants
Coalition partners have r = 0.42
Hamilton’s rule explains

why subordinates join the
coalition
rB = (0.42)(6.1) = 2.6 > 0.9 = C

Prediction of indirect selection favoring helping: helping should increase with
closer genetic relationship between potential helpers and young
-Females often breed synchronously; cubs are raised
together in a communal crèche
-Female lions feed their young and other young in
crèche
The proportion of nursing that goes to

non-offspring cubs increases with
relatedness of lionesses in group
Kin selection is important in cooperative
breeding of female lions
If indirect selection is an important mechanism in the evolution of altruism, we can
predict much of altruistic behavior is directed at genetic relatives
We find this to be the case in

many species, such as
Belding’s ground squirrels
What about cases of altruism toward non-relatives?

Non-breeding adults are “helpers,” as in
these meerkats
Cooperative breeding: adults other than

genetic parents help raise young
One dominant female and one
dominant male account for >
80% of group’s reproduction
Pups are cared for by the group
until 3 months
Helpers are young individuals who have not yet dispersed

Two major ways of helping:
Baby-sitting
Provisioning young with food
If helping is altruistic, there should be a benefit to others and a cost to the helper
Food intake and daily weight

gain of pups increases with
number of helpers
(benefit to others)
When litter size was manipulated,

helpers with high pup:helper ratios
(who had to help more) gained less
weight
(cost to helpers)
Prediction of indirect selection favoring helping: helping should increase with
closer genetic relationship between potential helpers and young
In meerkat groups, top baby-sitters spent 42% of

their day baby-sitting; bottom baby-sitters spent
only 8% of day baby-sitting
The helper that did the most baby-sitting was not more related
to young than the helper that did the least
Full siblings did not help more than others
In this system, relatedness does not explain variation in
helping behavior
Direct benefit of helping behavior in meerkats: predator detection
Individuals in larger groups spend less

time glancing around; interrupt foraging
less
Percentage of time at least one individual

is acting as sentinel generally increases
with group size
In meerkats, cooperative breeding likely evolved through

individual selection—helpers help to increase group size to
their own benefit
Cooperative breeding in the Florida Scrub Jay
-Mated pairs typically remain together for life, “divorce rate” is only 4%
-Live 10+ years
-RARE: both observational data and DNA indicate 0% infidelity
-Each year about half of all Florida Scrub-Jay territories are occupied by a
mated pair and 1 - 6 “helpers,”— usually comprised of offspring from previous
years.
-The longer a pair is together, the more helpers they have.
Benefit to the mated pair?

-Helpers increase the number of young raised each year.
-The breeding male is relieved of some of the responsibility of finding food for
the female and the young and can spend more time acting as a sentinel.
-Helpers care for the nestlings, search for food, and act as sentinels.
Benefit to the helpers?

-The many related helpers on a territory usually get a start on their own territory
by taking over a nearby territory, often aided by their parents
-Or by “budding off” a small territory adjacent to their parents.
•Vampire bats who fail to get a blood meal beg for food
Investigating altruism •Other bats may regurgitate blood for unsuccessful bat
•Donor & recipient bats are not always related
in non-relatives:
•Potential explanation: reciprocal altruism
vampire bats •Advantageous for individual A to help individual B if B later
reciprocates by helping A
What is the problem with
explaining this form of altruism?
•Reciprocal altruism may be unstable

in the face of cheating: selfish behavior
that evades altruistic conventions (i.e.
soliciting and taking help, failing to
reciprocate)
•Is there a way to protect a system of

reciprocity against cheating?
Successful strategies:
-Policing and punishing
-“Tit for Tat”
Tit for Tat: Begin with cooperation, then do whatever other player
did on previous move
Success of a strategy is based on being:
1) Retaliatory: punishing defection by retaliating

2) Forgiving: forgive after one retaliation so that
cooperation can be restored
3) Nice: never being the first to defect
Two major explanations for
altruism like cooperative
breeding:
Individual selection
-some aspects of cooperative
breeding have positive effects
on direct fitness of actor
-can be considered mutualism
Kin selection
-some aspects of cooperative breeding
can be explained by relatedness and
inclusive fitness of the actor

Altruism, Kin Selection, and Cooperative Breeding

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Altruism, Kin Selection, and Cooperative Breeding

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Altruism,

If an individual behaved in such a way that

These genes would not be passed on.

So, how can helping behavior evolve?

Nests are clumped in groups

If a male chases off a Bluegill sunfish

predator, this benefits

Bluegill sunfish males care

Nests are clumped in groups

Bluegill sunfish If a male eats the eggs in

Biological definition of altruism: by behaving

Belding’s ground squirrel’s “terrestrial predator” alarm call

Recipients of call benefit by being alerted to predator.

DIRECT FITNESS: a measure of an individual’s reproductive (genetic)

INDIRECT FITNESS: a measure of the genetic success of an altruistic

Inclusive fitness = a total measure of an individual's

Inclusive fitness = direct fitness + indirect fitness

r = the genetic relatedness of the recipient to the actor

C = the reproductive cost to the individual performing the act

Mom Dad Common coefficients (r)

Offspring Sibling Half sibs 1/4

Aunt to her nephew or

Dominant males have higher reproductive success

Coalition partners have r = 0.42

Hamilton’s rule explains

rB = (0.42)(6.1) = 2.6 > 0.9 = C

The proportion of nursing that goes to

We find this to be the case in

What about cases of altruism toward non-relatives?

Cooperative breeding: adults other than

Helpers are young individuals who have not yet dispersed

Food intake and daily weight

When litter size was manipulated,

In meerkat groups, top baby-sitters spent 42% of

Individuals in larger groups spend less

Percentage of time at least one individual

In meerkats, cooperative breeding likely evolved through

Benefit to the mated pair?

Benefit to the helpers?

•Reciprocal altruism may be unstable

•Is there a way to protect a system of

Success of a strategy is based on being:

1) Retaliatory: punishing defection by retaliating

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