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COOPERATION Tit-for-Tat and reciprocal altruism By-product mutualism We ’ ll save for later: Kin Selection Coop in Humans . Two roads to Cooperation. Coop. Defect. TFT. Defect. Coop. (b-c). -c. (-c). TFT. (b-c). Defect. b. 0. General payoff for costly
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COOPERATION • Tit-for-Tat and reciprocal • altruism • By-product mutualism • We’ll save for later: • Kin Selection • Coop in Humans
Two roads to Cooperation Coop Defect TFT Defect Coop (b-c) -c (-c) TFT (b-c) Defect b 0 General payoff for costly Cooperation Defect (b) 0 Prisoner's Dilemma: the only ESS is to Defect TFT is an ESS when: (-) c b > Reciprocal altruism OR Cooperation is an ESS when: r > c/b where r = coefficient of relatedness (Kin Selection)
Reciprocity – Food sharing in Vampire Bats (Gerald Wilkinson) Evidence for repeated interactions – unrelated group members may share a roost for several years
Data from natural roosts 1) Bats tend not to be related to each other 2) Cooperation occurs among commonly associating bats Associations Cooperating Pairs Associations All Pairs
Benefit of receiving aid outweighs the cost of donating cost of giving up 5% of your fed weight weight benefit of receiving that 5% starvation time since last meal (hrs)
Benefit of receiving aid outweighs the cost of donating cost of giving up 5% of your fed weight weight benefit of receiving that 5% starvation cost benefit time since last meal (hrs)
Using experimental roosts Wilkinson showed: -- 12 of 13 regurgitations were between members of the same roost in the field -- Recipients reciprocated to previous donors more than was expected by chance • probability of future interaction is high • individuals can recognize each other • the benefit of a cooperative act greatly outweighs the cost • potential for future retaliation is one does not cooperate
Egg trading and reciprocity in sequentially hermaphroditic fishes, e.g., black hamlet fish Switching gender roles as reciprocal altruism
Are these guppies cooperating? Predator-inspection behavior in guppies
Cooperate (inspect) Defect a b Cooperate Defect d c Yes, inspection is costly Payoff matrix in terms of the probability of not being eaten by the predator Is c > a ?
Information is Public This individual reacts similarly as the lcose individual when the latter is visible
Milinski’s experiments with mirrors and sticklebacks shows retaliation Mirror projects a fellow inspector traveling side by side the length of the aquarium Mirror projected 32° shows fellow inspector dropping back and finally disappearing the closer the focal fish approaches the predator
Reciprocal Altruism and predator mobbing
(1) Defector Cooperator (2)
Cooperate Defect 5 1 Cooperate Defect 0 1 Problems with Tit-for-Tat Byproduct mutualism Tracking debts owed and favors given requires complex score-keeping Cooperate is the sole ESS
Prisoner’s Dilemma and byproduct mutualism in blue jays • Trained blue jays choose a perch that • delivers: • Large rewards to its neighbor • Or • (2) Small rewards to itself
Prisoner’s Dilemma Byproduct Mutualism Prisoner’s Dilemma Mutual cooperation Mutual defection One coop, one defect ????? Why don’t they coop Here ???? • Individuals act in concert with the theory (in part) • Individuals learn quickly to respond accordingly • Individuals do not cooperate in the Prisoner’s Dilemma even when iterated Why the mutual defection even when the PD game is iterated between jays?
It is only through iterated cooperation that both jays can receive their largest payoff Individual play of the game has a large temptation to cheat – Prisoner’s Dilemma
The structure of the Payoff Matrix Matters – organisms learn/have knowledge of the payoffs and respond accordingly (Pris Dil vs. Byproduct Mut) Reciprocal Altruism (TFT) is broadly distributed in the animal world Its seen in simple organisms (fish) largely and social spp (humans later) Sex, social foraging, predator avoidance/mobbing, and others for sure
Accumulating benefits (low discount) Opponent’s strategy TFT Proportion cooperating Unaccumulating (high discount) The trick is for jays to see their food accumulating in transparent containers and every 4 turns they get to feed.