Differentiation is regulated at the genomic level by a preprogrammed sequence of gene activations and repressions. The main regulators are thought to be nonhistone chromosomal proteins, among which the DNA-binding proteins are the best candidates. However, histones and their variants most probably also play a role. In addition, it is conceivable that nuclear sap proteins and the nuclear skeleton, by temporarily interacting with the chromatin, might have accessory regulatory functions.
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Kuenzle et al. (1983) studied this question.