Kin Selection &

Inclusive Fitness

Social interaction
Social interactions between organisms present the opportunity
for conflict and cooperation
Performer benefits Performer suffers
Receiver benefits

Cooperation

Altruism

Receiver suffers

Selfish

Spiteful

Cooperation, or mutualism, results in fitness gain for both participants

Selfish actions benefit the performer and hurt the receiver
Spiteful actions hurt both parties (hurt oneself to hurt another)
Altruism, the performer suffers in order to benefit the recipient

Altruism

Behavior that benefits a receiver at a cost to the actor

Honey bee sting, Alarm calls, Blood sharing by bats
Cooperation: Displaying a behavior that benefits another individual.
(If both benefit that's mutualism)
Altruism: Displaying a behavior that benefits another individual at a
cost to oneself.
Social behavior is NOT cooperative behavior : it is possible to have
sociality with no cooperation and cooperation with no sociality.

http://eebweb.arizona.edu/Courses/Ecol600A/kin%20selection.pdf

Evolution of Altruism

How can Altruism evolve?

If the recipient of the altruistic act benefits, it is going to leave more
offspring.
The actor however is not going to leave more offspring, or even
fewer offspring fewer altruists in the next generation.
If such behavior is heritable, and it goes on over many generations,
it will ultimately die out.
Are there hidden benefits to helping somebody at a cost to yourself?

Group selection rare, only if long-term assortment maintained

Kin-selection yes, if helping relatives
Sexual selection yes, if mating benefits
Reciprocal altruism yes, if reciprocation likely and enforced
Status yes, if indirect benefits

http://eebweb.arizona.edu/Courses/Ecol600A/kin%20selection.pdf

Kin Selection

Helping relatives increases your inclusive fitness:

Inclusive fitness: your own offspring (fitness) plus
your genes reproduced in others.
This means that the more of your genes are in a
relative, the more interest you have in helping
them.
This is measured by r (relatedness), also called
coefficient of relationship
r is usually defined as the average proportion of
alleles of an individual A that are identical by
descent to those in individual B.
Or, the probability that A and B carry the same
allele, derived from the same ancestor, at a
particular locus.

http://eebweb.arizona.edu/Courses/Ecol600A/kin%20selection.pdf

Kin Selection
Hamiltons rule:
An individual can be altruistic if
c<b*r
The cost should be smaller than the benefit multiplied by
relatedness.
E.g. an individual may not reproduce in a given year (c=1) to help its
sibling (r=0.5) if this helps the sibling raise at least two additional
offspring (b=2).

Question: Would you lay down your life for

your brother?
No, but I would for
two brothers
or eight cousins
J.B.S. Haldane
http://eebweb.arizona.edu/Courses/Ecol600A/kin%20selection.pdf

Relatedness
Relatedness = 0.25

Relatedness = 0.5

If you have two

children, on average
one copy of each of
your genes survives.

http://eebweb.arizona.edu/Courses/Ecol600A/kin%20selection.pdf

If you have four

nieces, on average
one copy of each of
your genes survives.

This means, if you

sacrifice yourself for
four nieces, your
genes have lost
nothing.

Conditions for altruism towards genetic relatives (kin) set by Hamiltons rule:
b/c > 1/r
b = benefits to recipient
c = cost to donor
r = coefficient of relatedness
If r = 1/2, then benefit b must 2c.

Relationship

mother-offspring

1/2

sister-brother

1/2

uncle-nephew

1/4

cousin-cousin

1/8

Inclusive fitness

W.D. Hamilton (1964)

An individual increases its fitness either through own
personal reproduction OR reproduction of genetic relatives
Inclusive fitness is fitness due to both forms of reproduction

10_IB lecWEB_Papaj-1.ppt

Kin Selection

When it is demonstrated that individuals preferentially help kin rather

than non-kin, this is taken as evidence for kin-selection.
Examples for kin-selection:
social insects
prairie dog alarm calls to offspring & other relatives
generally parental care

Kin-recognition
Do individuals have to be able to recognize
relatives for kin selection to work?
NO kin selection can operate, and cause the
evolution of altruism, as long as altruists are
more likely to help kin than non-kin
- for whatever reason.

http://eebweb.arizona.edu/Courses/Ecol600A/kin%20selection.pdf

Males playing

Females fighting

Cooperation between male chimpanzees is greater than cooperation

between females.

In chimps, females disperse from natal troop, whereas males often stay.
Hence, males in a troop are more genetically-related to each other.

10_IB lecWEB_Papaj-1.ppt

Alarm calling: Beldings Ground Squirrel

Distance moved from

natal burrow (m)

Cooperation in chasing
trespassing squirrels

Highly social mammal; breeds in colonies

Females tend to breed near their birthplace
Neighbors are often closely related
Squirrels trill when predatory mammals are present and
whistle when hawks are seen
Individuals that whistle have lower mortality than those
that do not (selfish)
Individuals that trill have higher mortality than those that
do not (altruistic)

Age (months)
lecture 19-21 Kin Selection and social behavior.ppt

lecture 08 - kin selection + eusociality.ppt

Alarm calling: Beldings Ground Squirrel

Females were much

more likely to call
than males

Females were much more likely to

call when they had a close relative
nearby --thus, altruistic actions are
not randomly distributed

From Sherman et al. 1977

lecture 08 - kin selection + eusociality.ppt

Kin Selection in White-Fronted Bee-Eaters

In many birds, young will help their parents rear new offspring, rather than go
off and reproduce themselves
New offspring may be full or halfsiblings of the young helpers
Common in species where
opportunities for breeding or new
nest construction are limited
In these cases, the young helping
their parents may be making the
best of a bad situation
Bee-Eaters live in big colonies, where relatives and non-relatives nest in
close proximity, as part of the same clan (= nesting group)
1-yr olds often help at an existing nest instead of starting their own
Prediction: coeff. of relatedness should dictate whether birds help clan-mates
lecture 08 - kin selection + eusociality.ppt

Kin Selection in White-Fronted Bee-Eaters

(birds that havent yet reproduced)

(2) Young selectively help more closely related

adults (coefficient of relatedness predicts
how help will be distributed)

(1) Unrelated birds that have married

into a clan are much less likely to
help than a blood relative born into
the clan (= natal)

This makes a huge difference to parents

Half of bee-eater babies starve to death
before leaving the nest
Each 1-yr old helper results in 0.47 more
offspring being successfully raised
--major boost to inclusive fitness

Kin Selection

Inclusive fitness theory vs. kin selection

In fact, thats why some argue that it should be called inclusive
fitness theory rather than kin selection Altruism can evolve as
long as altruists are more likely than chance to dispense help to
other altruists.
Kin-recognition
By smell (rodents, humans, insects)
By song (some birds)
By learning/familiarity (mice, humans)
By visual similarity (chimpanzees, humans)

http://eebweb.arizona.edu/Courses/Ecol600A/kin%20selection.pdf

Kin Selection & Eusociality

Eusociality: some individuals sterile (caste)

Evolved > 10 times in Hymenoptera (haplodiploid)
All members of a colony are usually highly related
Haplodiploidy: sex determining mechanism in which
males are haploid (1N) and females diploid (2N).
Haplodiploidy cause relatedness asymmetry
Does haplodiploidy cause eusociality?
In complete monogamy, workers are more related to
the queens daughters (sisters) (r=0.75) than to their
own (r=0.5)
Worker bees can benefit by foregoing reproduction and
helping queen rear more sisters
This would explain why so many Hymenoptera are
eusocial and why workers are always females

http://eebweb.arizona.edu/Courses/Ecol600A/kin%20selection.pdf

Kin Selection & Eusociality

However, workers are only related to

males by r=0.25 (less than to
daughters) thus average relatedness
to reproductive offspring is still 0.5
(depending on sex ratio)
Actual relatedness values measured in
insect colonies are almost never 0.75
(multiple queens, polygamy)
Some eusocial species lack
haplodiploidy
Sterile Castes in Naked Mole Rats
Occur in colonies of up to 300 members.
Breeding restricted to a single queen.
Inbreeding results in high relatedness

Many haplodiploid species are not

social

http://eebweb.arizona.edu/Courses/Ecol600A/kin%20selection.pdf

Facultative Eusociality
Totipotency of only the more
reproductive caste
Multi-foundress nests grew fastest
when there was a big difference in
body size between the dominant
queen and her subordinates
-less time lost to challenges to the
dominant queen by her underlings
(fighting for the right to lay more
eggs)
Subordinates may help instead of starting your own nest because
subordinates are close relatives of the dominant foundress

Facultative Eusociality
Possible strategies:
-strike out on your own, risk it
-help a close relative, thereby increasing your inclusive fitness
-sit and wait: dont help anyone, but wait for chance to steal a
nest should a foundress die or you can beat her up later
-leave nest in early spring, hibernate, try again next season
-reproductive altruism is facultative, meaning it can be adopted
depending on the conditions faced by a particular female
-relatedness
-body size
-nest availability
-time of year (season)

Eusociality

Alternative hypotheses for the origin of eusociality:

Parental manipulation
Predisposition to sociality because of high b/c ratio
(underground nests, extended brood care)
Group selection

Wilson & Hlldobler 2005

Superiority of colony life over solitary life (b may be much greater than c)
Eusociality arose among unrelated individuals first; then relatedness increased
In many species nests are founded by unrelated individuals
Real-existing relatedness low and counterproductive (?)
Eusociality rare even in highly related groups

Conclusions from the controversy

Haplodiploidy is not crucial to evolution of eusociality
Ecological factors (high b/c) explain most of the variation between species in sociality
Controversy arises over the definition of r relatedness by pedigree or measure of
genetic similarity?
Complete worker sterility can only arise with positive r, whether by kinship or other
segregation mechanisms
However, many social insects do not actually have complete worker sterility

http://eebweb.arizona.edu/Courses/Ecol600A/kin%20selection.pdf

Kin Selection and Parental Care

Predictions of kin selection about parental care

Individuals should invest in (care for) their own offspring versus
unrelated young

Kin selection in Bluegill

http://instruct.uwo.ca/zoology/336a/lecture4handout.pdf

Hain and Neff 2006

Kin Selection and Mating System

Within nests, the average relatedness of parentals offspring is >3x

that of cuckolders offspring
Thus, a parentals offspring could gain more kin-selective benefits
than a cuckolders offspring simply by associating with and helping a
random nestmate
Kin discrimination may be important when there is an optimal group
size that limits membership or when food resources are limited and
shared among group members.
But, to gain a similar kin-selective
benefit, a cuckolders offspring would
have to actively discriminate among
nestmates, which they do via a selfreferent odor mechanism
Experimental evidence from
behavioral trial that separate kin
recognition from familiarity
Hain and Neff 2006

Reciprocal altruism

Altruistic behavior among unrelated individuals

Trivers (1971) hypothesized that individuals may be selected to
dispense altruistic acts if equally-valuable favors are returned in the
future (Reciprocal altruism)
May evolve if:
The cost to the actor is less than or equal to the benefit to the recipient
Individuals that do not reciprocate are punished

May evolve when:

Interactions generally take place among the same sets of individuals

Many opportunities for altruistic behavior exist
Individuals have good memories
Costs vs. benefits are roughly equal

Blood sharing in Vampire Bats

Roost in hollow trees

High levels of association in roosting behavior
Regurgitate blood meals to one another: 33% of young bats
and 7% of adults fail to feed on any given night
Kin selection or reciprocal altruism?

Blood sharing in Vampire Bats

Five criteria to distinguish reciprocity vs kin selection (Wilkinson 1990):
1: Females associate for long periods, so there are numerous chances to be
either the donor or recipient bat.
2: The likelihood of regurgitation to roostmates can be predicted on the
basis of past associations.
3: The roles of donor and recipient reverse often.
4: The short-term benefits to the recipient outweigh the costs to the donor.
5: Donors can recognize and expel cheaters to this system.

Kin selection or reciprocal altruism? Probably a little of both

(most sharing between mother-pups, but also between unrelated buddies)