Model for coordination of the meiotic program by Cdc55. (A) In wild-type cells during metaphase I, PP2ACdc55 both maintains Cdc14 sequestration in the nucleolus by dephosphorylating Cfi1 and restricts the amount of PP2A scaffold (A) and catalytic (C) subunits available to form a complex with Rts1 (B′). The level of PP2ARts1 defines the domain of cohesion that will be protected during meiosis I. Once chromosomes are correctly aligned, separase activation leads to both the cleavage of unprotected arm cohesin and, through PP2ACdc55 down-regulation, Cdc14 release. Cdc14 release during anaphase I ensures that a second spindle axis will assemble in meiosis II after its resequestration. (B) In cdc55mn cells, ectopic Cdc14 release interferes with spindle assembly, and excess PP2ARts1 extends the domain of protected cohesin. This prevents nuclear division except in a low number of cells in which Cdc14 is relocalized to the nucleolus through an unknown mechanism (?). P indicates Cdk-dependent phosphorylation of Cfi1/Net1.