In the fission yeast Schizosaccharomyces pombe, the proteins Cut1 and Cut2 form a complex that is catalytically inactive. At the onset of anaphase, Cut2 is polyubiquitylated by a large E3 complex containing Cut4, Cut9, Cut23, and other proteins, and is subsequently destroyed. Cut1 then cleaves Rad21, a non-SMC component of a complex that also contains two SMC proteins, thus allowing sister-chromatid separation. Mutations in the cut genes lead to the cut phenotype, in which the cell attempts cytokinesis without chromosome segregation. Indicate which of the following proteins or protein complexes corresponds to or contains the product of the genes cut1 (A), cut2 (B), cut4 (C), and rad21 (D). Your answer would be a four-letter string composed of letters A to D only, e.g. DCAB. ( ) Securin ( ) Cohesin ( ) Anaphase-promoting complex ( ) Separase