178 Heterochromatin: new possibilities for the inheritance of structure Shiv IS Grewal* and Sarah CR Elgin† Significant portions of the eukaryotic genome are Two key observations have linked formation of such a heterochromatic, made up largely of repetitious sequences and condensed heterochromatic structure with the inactivation possessing a distinctive chromatin structure associated with of genes normally resident in euchromatic domains. First, gene silencing. New insights into the form of packaging, the X chromosome inactivation in mammals leaves the associated histone modifications, and the associated inactive X as a visibly staining structure, the Barr body. nonhistone chromosomal proteins of heterochromatin have Although the choice of which chromosome to inactivate — suggested a mechanism for providing an epigenetic mark that either maternal or paternal — appears to be random in allows this distinctive chromatin structure to be maintained most mammalian species, the decision is clonally inherited following replication and to spread within a given domain. once made [4]. Second, in Drosophila, a similar phenome- non of clonally inherited silencing is observed following Addresses chromosome rearrangements with one breakpoint within *Cold Spring Harbor Laboratory, One Bungtown Road, heterochromatin (position effect variegation [PEV]; see Cold Spring Harbor, New York 11724, USA; Figure 1). For example, juxtaposition of the white gene e-mail:
[email protected] with such a breakpoint results in silencing of white in some †Washington University, One Brookings Drive, Department of Biology, CB-1229, St Louis, Missouri 63130, USA; of the cells in which the gene is normally active; patches of e-mail:
[email protected] expressing cells are observed, again suggesting a stochastic ‘decision’ stably inherited through mitosis.