Inclusive fitness and the major transitions in evolution

Inclusive fitness theory is the leading framework for explaining the major transitions in evolution, whereby free-living subunits (e.g. cells, organisms) have cooperated to form new, higher-level units (e.g. organisms, eusocial societies). The theory has attracted considerable controversy. From a brief survey of the controversy's present status, I conclude that inclusive fitness theory continues to provide both a concept and a principled modelling tool of value for understanding social evolution, including major transitions. Turning to new developments in the study of major transitions, I describe work defining the point of occurrence of major transitions and, from inclusive fitness theory, the required conditions. I also suggest that it remains important to understand the evolution of individuality that occurs beyond such thresholds.