Centromeres fall apart (right) without Hsk1 action in S phase.
Forsburg/Macmillan
The dual action makes sense, as chromosomes must be stuck together as soon as they are replicated. The responsible kinase activity, Hsk1 (CDC7)–Dfp1, is restricted to S phase, when DNA replication takes place.
Dfp1 turned up in a two-hybrid screen with Swi6, the fission yeast equivalent of heterochromatin protein 1 (HP1). Cells with a mutant Dfp1 that no longer binds Swi6 can replicate their DNA but suffer segregation errors when their defective centromeres fall apart. Swi6 localization is normal in these cells but, based on in vitro results, Swi6 phosphorylation may be reduced. This is the first indication that Swi6 localization is not sufficient to define heterochromatin function.
An interesting parallel is known in budding yeast, where establishment of silent heterochromatin at the mating type locus requires passage through S phase, though not DNA replication. Budding yeast lacks an HP1 homologue, but perhaps other proteins serve an equivalent heterochromatic function. ▪
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