Shugoshins: from protectors of cohesion to versatile adaptors at the centromere.

Sister chromatids are held together by a protein complex named cohesin. Shugoshin proteins protect cohesin from cleavage by separase during meiosis I in eukaryotes and from phosphorylation-mediated removal during mitosis in vertebrates. This protection is crucial for chromosome segregation during mitosis and meiosis. Mechanistically, shugoshins shield cohesin by forming a complex with the phosphatase PP2A, which dephosphorylates cohesin, leading to its retention at centromeres during the onset of meiotic anaphase and vertebrate mitotic prophase I. In addition to this canonical function, shugoshins have evolved novel, species-specific cellular functions, the mechanisms of which remain a subject of intense debate, but are likely to involve spatio-temporally coordinated interactions with the chromosome passenger complex, the spindle checkpoint and the anaphase promoting complex. Here, we compare and contrast these remarkable features of shugoshins in model organisms and humans.