Musacchio, A. The molecular biology of spindle assembly checkpoint signaling dynamics. Curr. Biol. 25, R1002–R1018 (2015).
Nasmyth, K. & Haering, C. H. Cohesin: its roles and mechanisms. Annu. Rev. Genet. 43, 525–558 (2009).
Uhlmann, F., Wernic, D., Poupart, M. A., Koonin, E. V. & Nasmyth, K. Cleavage of cohesin by the CD clan protease separin triggers anaphase in yeast. Cell 103, 375–386 (2000).
Mei, J., Huang, X. & Zhang, P. Securin is not required for cellular viability, but is required for normal growth of mouse embryonic fibroblasts. Curr Biol 11, 1197–1201 (2001).
Pfleghaar, K., Heubes, S., Cox, J., Stemmann, O. & Speicher, M. R. Securin is not required for chromosomal stability in human cells. PLoS Biol. 3, e416 (2005).
Llano, E. et al. Shugoshin-2 is essential for the completion of meiosis but not for mitotic cell division in mice. Genes Dev. 22, 2400–2413 (2008).
Orth, M. et al. Shugoshin is a Mad1/Cdc20-like interactor of Mad2. EMBO J. 30, 2868–2880 (2011).
Boland, A. et al. Cryo-EM structure of a metazoan separase-securin complex at near-atomic resolution. Nat. Struct. Mol. Biol. 24, 414–418 (2017).
Lin, Z., Luo, X. & Yu, H. Structural basis of cohesin cleavage by separase. Nature 532, 131–134 (2016).
Wirth, K. G. et al. Separase: a universal trigger for sister chromatid disjunction but not chromosome cycle progression. J. Cell. Biol. 172, 847–860 (2006).
Zou, H., McGarry, T. J., Bernal, T. & Kirschner, M. W. Identification of a vertebrate sister-chromatid separation inhibitor involved in transformation and tumorigenesis. Science 285, 418–422 (1999).
Boos, D., Kuffer, C., Lenobel, R., Korner, R. & Stemmann, O. Phosphorylation-dependent binding of cyclin B1 to a Cdc6-like domain of human separase. J. Biol. Chem. 283, 816–823 (2008).
Hellmuth, S. et al. Positive and negative regulation of vertebrate separase by Cdk1-cyclin B1 may explain why securin is dispensable. J. Biol. Chem. 290, 8002–8010 (2015).
Huang, X. et al. Preimplantation mouse embryos depend on inhibitory phosphorylation of separase to prevent chromosome missegregation. Mol. Cell. Biol. 29, 1498–1505 (2009).
Huang, X. et al. Inhibitory phosphorylation of separase is essential for genome stability and viability of murine embryonic germ cells. PLoS Biol. 6, e15 (2008).
Huang, X., Hatcher, R., York, J. P. & Zhang, P. Securin and separase phosphorylation act redundantly to maintain sister chromatid cohesion in mammalian cells. Mol. Biol. Cell 16, 4725–4732 (2005).
Stemmann, O., Zou, H., Gerber, S. A., Gygi, S. P. & Kirschner, M. W. Dual inhibition of sister chromatid separation at metaphase. Cell 107, 715–726 (2001).
Li, M., York, J. P. & Zhang, P. Loss of Cdc20 causes a securin-dependent metaphase arrest in two-cell mouse embryos. Mol. Cell. Biol. 27, 3481–3488 (2007).
Hellmuth, S. et al. Human chromosome segregation involves multi-layered regulation of separase by the peptidyl-prolyl-isomerase Pin1. Mol. Cell 58, 495–506 (2015).
Gorr, I. H., Boos, D. & Stemmann, O. Mutual inhibition of separase and Cdk1 by two-step complex formation. Mol. Cell 19, 135–141 (2005).
Rodriguez-Bravo, V. et al. Nuclear pores protect genome integrity by assembling a premitotic and Mad1-dependent anaphase inhibitor. Cell 156, 1017–1031 (2014).
Holland, A. J. & Taylor, S. S. Cyclin-B1-mediated inhibition of excess separase is required for timely chromosome disjunction. J. Cell Sci. 119, 3325–3336 (2006).
Clift, D., Bizzari, F. & Marston, A. L. Shugoshin prevents cohesin cleavage by PP2A(Cdc55)-dependent inhibition of separase. Genes Dev. 23, 766–780 (2009).
Tsou, M. F. & Stearns, T. Mechanism limiting centrosome duplication to once per cell cycle. Nature 442, 947–951 (2006).
Kawashima, S. A., Yamagishi, Y., Honda, T., Ishiguro, K. & Watanabe, Y. Phosphorylation of H2A by Bub1 prevents chromosomal instability through localizing shugoshin. Science 327, 172–177 (2010).
Hauf, S. et al. Dissociation of cohesin from chromosome arms and loss of arm cohesion during early mitosis depends on phosphorylation of SA2. PLoS Biol. 3, e69 (2005).
Eytan, E. et al. Disassembly of mitotic checkpoint complexes by the joint action of the AAA-ATPase TRIP13 and p31comet. Proc. Natl Acad. Sci. USA 111, 12019–12024 (2014).
Reddy, S. K., Rape, M., Margansky, W. A. & Kirschner, M. W. Ubiquitination by the anaphase-promoting complex drives spindle checkpoint inactivation. Nature 446, 921–925 (2007).
Bakos, G. et al. An E2-ubiquitin thioester-driven approach to identify substrates modified with ubiquitin and ubiquitin-like molecules. Nat. Commun. 9, 4776 (2018).
Hellmuth, S., Böttger, F., Pan, C., Mann, M. & Stemmann, O. PP2A delays APC/C-dependent degradation of separase-associated but not free securin. EMBO J. 33, 1134–1147 (2014).
Gorr, I. H. et al. Essential CDK1-inhibitory role for separase during meiosis I in vertebrate oocytes. Nat. Cell Biol. 8, 1035–1037 (2006).
Hellmuth, S., Gutiérrez-Caballero, C., Llano, E., Pendás, A. M. & Stemmann, O. Local activation of mammalian separase in interphase promotes double-strand break repair and prevents oncogenic transformation. EMBO J. 37, e99184 (2018).
Schöckel, L., Möckel, M., Mayer, B., Boos, D. & Stemmann, O. Cleavage of cohesin rings coordinates the separation of centrioles and chromatids. Nat. Cell Biol. 13, 966–972 (2011).
Murray, A. W. Cell cycle extracts. Methods Cell Biol. 36, 581–605 (1991).
McGuinness, B. E., Hirota, T., Kudo, N. R., Peters, J. M. & Nasmyth, K. Shugoshin prevents dissociation of cohesin from centromeres during mitosis in vertebrate cells. PLoS Biol. 3, e86 (2005).
Franken, N. A., Rodermond, H. M., Stap, J., Haveman, J. & van Bree, C. Clonogenic assay of cells in vitro. Nat. Protocols 1, 2315–2319 (2006).
Butt, T. R., Edavettal, S. C., Hall, J. P. & Mattern, M. R. SUMO fusion technology for difficult-to-express proteins. Protein Expr. Purif. 43, 1–9 (2005).
Holland, A. J., Bottger, F., Stemmann, O. & Taylor, S. S. Protein phosphatase 2A and separase form a complex regulated by separase autocleavage. J. Biol. Chem. 282, 24623–24632 (2007).
Schägger, H. Tricine-SDS-PAGE. Nat. Protocols 1, 16–22 (2006).