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Cooperation
Cooperation is an outcome in which two or more individuals interact in a way that benefits all participants, even if they each incur some cost.
Two meaning of cooperate:
achieving teamwork within a group
Individual behaviors that enable the group to succeed
Benefits of cooperation
Why animals (or humans) cooperate:
Access to food and water
Protection and reduced aggression
Lower tension or stress
Coalition formation (social bonds)
Group acceptance (“team player” advantage)
Mating opportunities
Examples of cooperative behavior
Male-male alliances (stallions):
subordinate males from alliances with dominant males
both benefit through increased access to females and reduced conflict
Social grooming (allogrooming):
one individual grooms another
common in primates, horses, and mammals- builds bonds and hygiene
Symbiotic relationships:
example: cattle egrets and horses- birds eat pests off large mammals
both species benefit → mutualistic cooperation
The four paths to cooperation
There are four major evolutionary pathways to cooperation:
Reciprocity- exchange of favors
Mutualism- immediate benefits for all participants (no cheating temptation)
Kin selection- helping relatives to ensure gene survival
Group Selection- cooperation benefits the entire group, making it more successful than others
Reciprocity (exchange of favors)
Reciprocal altruism: helping unrelated individuals with the expectation of future help
Key mechanism: game theory- a mathematical model of strategic interaction.
Used to explain cooperation through prisoner’s dilemma and tit- for- tat strategies
The prisoner’s dilemma
Scenario: two prisoners are arrested. They can either cooperate (stay silent) or defect (betray each other)
Outcome:
Both cooperate → mild punishment (2 years) → mild punishment
One defects → betrayer goes free → partner gets 5 years
Both defect → both gets 3 years → both lose trust
Key insight:
even though cooperation yields the best group results, defection seems safer individually- creating a dilemma.
Game variables
T- temptation to cheat
R- reward for mutual cooperation
P- punishment for mutual defection
S- sucker’s payoff (for cooperating alone)
Must satisfy: T>R>P>S
Evolutionary stable strategy (ESS)
an ESS is a behavioral strategy that cannot be outcompeted once adapted by most of the population.
Axelrod & Hamilton used simulations to find that Tit- for- tat (TFT) is an effective ESS when players repeatedly meet
Tit-For-Tat (TFT) strategy
Rules of TFT:
Nice- cooperate first
Retaliatory- defect when the partner cheats
Forgiving- return to cooperation after the partner does
Neurobiology of Cooperation
Human experiments (using fMRI and PET scans) during Prisioner’s dilemma games show:
Brain activity increases during mutual cooperation
Cooperation activates reward centers → emotionally satisfying
Neuroeconomics studies how trust, empathy, and fairness influence decision making
Mutualism (Byproduct Cooperation)
Definition: cooperation where everyone benefits immediately, and there’s no temptation to cheat
Difference from reciprocity:
no need to track past interactions or punish defectors
Conclusion: learning and immediate benefits encourage cooperation better than delayed reciprocity
Group Selection (trait group selection model)
Definition: natural selection acts both within groups and between groups
Within group selection → favors selfish individuals who avoid cost of cooperation→ reduces coorperation
Between group selections → favors cooperative groups that outcompete selfish ones → promotes cooperation
Coalitions and Alliances
Coalition: cooperative action by two or more individuals/ groups against another
Alliance: long term coalition
Phylogeny and Cooperation
Phylogeny: evolutionary history shows traits inherited from common ancestors
Cooperative behavior may stem from shared ancestry, not independent evolution.
Conclusion: some cooperation traits evolved once and were passed down through lineages
Interspecific Cooperation (Mutualism)
Intraspecific: cooperation within a species
Interspecific: cooperation between species- mutualism