This chapter tackles the evolution of cellular organization. How did intricate subcellular machines, such as ribosomes, flagella and ion pumps come to exist? How do cells transmit their functional organization to their offspring, and how did that evolve? And where did cellular organization come from in the first place? Contrary to the claims of Intelligent Design, there is ample evidence that random variation of genes winnowed by natural selection played a large role. But conventional views on these matters are too restrictive. Cells transmit structural organization by a hierarchy of mechanisms that includes genes, self-organization, the continuity of membranes and other structures, and a role for the cytoskeleton in helping a growing cell to model new structures upon the existing ones. How cells as we know them originated remains to be discovered, a subject for speculation and wonder but not yet for explication.