Cell Science at a Glance 4139 The spindle checkpoint mechanism that delays anaphase onset highlight current understanding of how until all chromosomes are correctly the spindle checkpoint is activated, how it Karen M. May and Kevin G. attached in a bipolar fashion to the delays anaphase onset, and how it is Hardwick mitotic spindle. silenced. Wellcome Trust Centre for Cell Biology, University of Edinburgh, EH9 3JR, UK The core spindle checkpoint proteins are (e-mail:
[email protected]) Mad1, Mad2, BubR1 (Mad3 in yeast), Activation of the checkpoint Journal of Cell Science 119, 4139-4142 Bub1, Bub3 and Mps1. The Mad and Bub During mitosis spindle microtubules Published by The Company of Biologists 2006 proteins were first identified in budding bind to complex protein structures called doi:10.1242/jcs.03165 yeast by genetic screens for mutants that kinetochores, which assemble on the failed to arrest in mitosis when the spindle centromere of each chromosome. The Every mitosis, replicated chromosomes was destroyed (Taylor et al., 2004). These Mad and Bub proteins localise to the must be accurately segregated into each proteins are conserved in all eukaryotes. outer kinetochore early in mitosis, before daughter cell. Pairs of sister chromatids Several other checkpoint components, proper attachments are established, and attach to the bipolar mitotic spindle such as Rod, Zw10 and CENP-E, have accumulate on unattached kinetochores. during prometaphase, they are aligned at since been identified in higher eukaryotes When spindle microtubules make metaphase, then sisters separate and but have no yeast orthologues (Karess, contact with the outer kinetochore are pulled to opposite poles during 2005; Mao et al., 2003).