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what is cooperation?
a behaviour that benefits another individual and also increases the actor’s inclusive fitness (directly or indirectly)
what is true altruism?
a behaviour that reduces the actor’s inclusive fitness (direct + indirect), very rare and generally selected against
what is selfishness?
behaviour that increases the actor’s fitness at the expense of others
what is spite?
behaviour that reduces the actor’s fitness and also reduces the recipient’s fitness, extremely rare in nature
what is direct fitness?
fitness gained through producing one’s own offspring
what is indirect fitness?
fitness gained through helping relatives reproduce, weighted by relatedness (r)
what is inclusive fitness?
direct fitness + indirect fitness
selection acts on inclusive fitness, explaining helping behaviours toward kin
what is kin selection?
natural selection favouring helping behaviours directed at relatives, because they share alleles
helping occurs when rb > c (Hamilton’s rule)
what is reciprocal altruism?
cooperation between non-relatives where help is given with expectation of future return
requires repeated interactions, memory, and ability to punish cheaters
degrees of relatedness
parent to offspring - r = 0.5
full siblings - r = 0.5
half-siblings - r = 0.25
first-cousins - r = 0.125
grandparent to grandchildren - r = 0.25
what is Hamilton’s rule?
helping behaviour is favoured when: rb > c
r = relatedness to the recipient
b = benefit to recipient
c = cost to actor
reformulated Hamilton’s rule (when considering different viewpoints)?
helping is favoured when: r1b > r2c
r1 = actor’s relatedness to recipient
r2 = actor’s relatedness to the helper (themselves or someone else’s cost)
why is helping descendants not “altruism”?
because helping offspring increases direct fitness (passing on alleles), it is cooperation, not altruism
example: should you sacrifice yourself to save 5 cousins (r=0.125) and 2 half siblings (r=0.25)?
calculate rb:
(5 × 0.125) + (2 × 0.25) = 1.125 > 1 life cost → yes, kin selection favours sacrifice
example: should a bird help parents raise 4 full siblings or reproduce itself, producing 3 offspring?
helping siblings: rb = 0.5 × 4 = 2
reproducing: rb = 0.5 × 3 = 1.5
help siblings (higher inclusive fitness)
helping half-sister increases her fitness by 20 units. Mother’s POV: max cost you should pay?
mother’s relatedness to both of you = 0.5
her rule = 0.5(20) > 0.5c → 10 > c
c < 10
correct answer: c. help only if c < 10 units
father’s POV with the same problem?
father only shares genes with you, not your half sister
so r2 > r1 → no benefit
correct answer: no nonzero cost is acceptable
stepfather’s POV?
he shares no genes with either child, so he is indifferent
correct: always acceptable (no genetic cost/benefit)
why is there conflict even among close relatives?
genetic interests overlap but are not identical (ex: siblings value their own reproduction more than a sibling’s)
how do asymmetries in relatedness generate conflict?
if relatives differ in r-value to each other and to themselves, optimal decisions for one relative may differ from optimal decisions for another
what game model explains cooperation vs cheating between non-relatives?
the prisoner’s dilemma, where typical payoffs
T > R > P > S
T = temptation to cheat
R = reward for mutual cooperation
P = punishment for mutual cheating
S = sucker’s payoff (you cooperate, partner cheats)
why is cheating the “rational” one-shot choice?
because T > R and P > S
no future consequences → cheating maximizes fitness
under what conditions can cooperation evolve among unrelated individuals?
when interactions are repeated and individuals can:
recognize patners
remember past behaviour
reward helpers
punish/avoid cheaters
found in wolves, chimps, humans
why are “social emotions” useful in evolution?
emotions like guilt, trust, resentment maintain social scorekeeping, promoting cooperation and detering cheating