Evolutionary transitions to new levels of individuality have usually been treated as a part of the "units of selection" problem. It has previously been assumed that the unit of transmission and heritable variation at each level is the same--that it is the DNA base sequence and its variations. It is suggested here that considering the nature and the role of hereditary variations produced by non-DNA inheritance systems is essential for understanding some evolutionary transitions. The argument is illustrated by considering the role of epigenetic inheritance systems (EISs), the systems responsible for cellular memory, in the transition from unicellularity to multicellularity. It is argued that EISs played a vital role in the transition to multicellularity and the evolution of complex ontogenies, as well as having an important effect on the evolution of developmental strategies which protect the multicellular individual from disintegrating into its component parts. An analogy between the transition to multicellularity and the transition to a cultural unit integrated by language is also suggested.
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