Multilevel Selection

Price Equation: Δz̄ = Cov(w,z)/w̄ + E(w·Δz)/w̄

Parameters

Price Equation Decomposition

Global cooperators:
Between-group sel:
Within-group sel:
Generation:0
Between-group: altruist groups have higher fitness
Within-group: defectors free-ride on cooperators
Cooperation survives when between-group selection outweighs within-group selection