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Levels-of-selection analyses usually focus on particles nested within a collective (a), for example, individuals within a society, or cells within an organism. In the case of proto-mitochondria within a proto-eukaryote, much of the collective is not contained within the particles, as emphasized in (b) with the cytosol shaded. The cytosol can be viewed as an emergent feature of the collective. The host genome appears as the unique particle.
