Lect.

17: Social behavior Altruism: Cooperative male

display, alarm calling
• Altruism & cooperation
• Exam II: 1 Nov. Study guide
• Inclusive fitness, kin selection
Tuesday, 24 Oct
– Hamilton’s rule
• Seminar, 4:10, Today, 306 Abelson
– Relatedness

• Eusociality and Hymenopterma – Bethany Marshall: “Forensic

entomology”
– Sex ratio bias

• Eusociality in diploids

A solution? Kin selection

Hamilton’s rule
JBS: “If I had a one in 10 Favors traits that increase rB - C > 0
chance of drowning but
saved the life of my child, I indirect fitness WD Hamilton 1964

would save 5 copies of Natural Selection favors altrusistic acts

genes for this when indirect fitness benefits to the
behaviour….” JBS Haldane,
1953 receiver, reduced by the coefficient of
Inclusive Direct Indirect relatedness, exceeds costs to the
Fitness Fitness Fitness
altruist

Hamilton’s Rule
Who is the altruist?

• r, coefficient of Altruism
altruism relatedness
– Probability that
homologous alleles in
two individuals are
IBD Cost to actor (C) Benefit to recipient (B)
Individual reproduction Kin’s additional C = 1 offspring B = 2 offspring
reproduction
rB-C > 0
 r = coefficient of relatedness r = coefficient of relatedness r = coefficient of relatedness
= probability that an allele found in one
Relationship r
individual will be found also in another due to
inheritance from common ancestor (IBD). Parent - offspring 0.5
Full siblings 0.5
Relationship r Relationship r
Half siblings 0.25
P - O 0.5 full sib 0.5*0.5 + 0.5*0.5 Cousins 0.125
= 0.50
0.5 0.5
half sib 0.5*0.5 = 0.25
A A
Belding’s Ground Squirrel

BGS: “I will alarm call Who alarm calls?

and risk my life for 2 Females call for terrestrial predators
siblings, 4 half-sibs, or • BGS dispersal: Male >
8 cousins” female

rB - C > 0?

rB - C > 0? Male Avg. RS Male Avg. RS

Solo 0.9 young Solo 0.9 young
Dom. 7.0 young Dom. 7.0 young
Subord. 0.0 young Subord. 0.0 young

Cost = reduced # of offspring to subord. Benefit = # extra offspring to dominant

= 0.9 - 0.0 = 0.9 young = 7.0 - 0.9 = 6.1 young

Krakauer (2005) Nature 434:69-72 Krakauer (2005) Nature 434:69-72 Krakauer (2005) Nature 434:69-72
 Solve for relatedness? Cooperative mating: are male Inclusive fitness?
B = 6.1
coalitions related?
C = 0.9 B = 6.1 young
C = 0.9 young
rB - C > 0 r = 0.42

Ownbyphotography.com
Altruism pays: Net gain:
r > C/B rB-C > 0
> 0.9/6.1 1.7 > 0
> 0.15
Krakauer (2005) Nature 434:69-72 Krakauer (2005) Nature 434:69-72

Eusociality: reproductive How does eusociality evolve? Eusociality & hymenoptera

altruism • In Hymenoptera?
Sisters share 100% the genes they inherit from
– Haplodiploid sex determination:
• Specialized their father, which is 50% of their genome
– fertilized eggs (diploid) -> females
reproductive caste
– unfertilized eggs (haploid) -> males.
(Queen) Relationship r
WD Hamilton 1972
• Non-reproductive Q Q
castes (e.g. workers, fem - mom 0.5
drones, soldiers)
sis - bro 0.5*0.5 = 0.25
• Common in the reproductive sterile worker soldier

Hymenoptera: bees, Q W W W
W W sis - sis 0.5*0.5 +
wasps, ants
0.5*1 = 0.75

Kin selection & Hymenoptera? Fisher’s sex ratio theory Kin selection & Hymenoptera?
• Favors altruistic Males < females Males > females
• Favors biased sex ratio
female worker caste in
haplodiploids
females
Fitness

– Helping to raise their • Frequency- equilibrium Worker-Worker r = 0.75

sisters (r = 0.75)
dependent Worker-Brother r = 0.25
males
• Favors biased sex fitness Predicted sex ratio 75:25
ratio reproductive sterile worker soldier • Predicts a
– Workers raise offspring
50:50 sex ratio
W 0 50 100
% males in population
 Haplodiploidy & Eusociality? Haplodiploidy & Eusociality? How does eusociality evolve?
0.8
siblings
• But, multiple mating by queen erodes • Haplodiploidy?

Relatedness
0.6
relatedness among sisters – Not all haplodiploids are
• Multiple mating by eusocial
Diploid sibs
queen erodes – Eusocial Diploids
0.4
relatedness among females • Termites
• A shrimp
sisters 0.2
half-sibs • Naked mole rats

0.0 Genetic & Ecological

1 3 5 7 9
factors favor eusociality
Number of mates

Kin selection & Social behavior

• Mechanisms to explain altruism
• Hamilton’s rule: rB - C > 0
• Benefits of altruistic act discounted by r
• Relatedness in haplodiploid systems helps
explain eusociality and biased sex ratios
• Reproductive altruism favored by
– Genetic factors: Haplodiploidy, inbreeding
– Ecological factors