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Vol. 15, Issue 7, 3345-3356, July 2004

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Disruption of Astral Microtubule Contact with the Cell Cortex Activates a Bub1, Bub3, and Mad3-dependent Checkpoint in Fission Yeast

Sylvie Tournier ^*  , Yannick Gachet  , Vicky Buck ^*, Jeremy S. Hyams  , and Jonathan B.A. Millar ^* ||

^* Division of Yeast Genetics, National Institute for Medical Research, London NW7 1AA, United Kingdom; Department of Biology, University College London, London, WC1E 6B, United Kingdom

Submitted March 26, 2004; Accepted April 29, 2004
Monitoring Editor: Mark Solomon

In animal and yeast cells, the mitotic spindle is aligned perpendicularly to the axis of cell division. This ensures that sister chromatids are separated to opposite sides of the cytokinetic actomyosin ring. In fission yeast, spindle rotation is dependent upon the interaction of astral microtubules with the cortical actin cytoskeleton. In this article, we show that addition of Latrunculin A, which prevents spindle rotation, delays the separation of sister chromatids and anaphase promoting complex-mediated destruction of spindle-associated Securin and Cyclin B. Moreover, we find that whereas sister kinetochore pairs normally congress to the spindle midzone before anaphase onset, this congression is disrupted when astral microtubule contact with the actin cytoskeleton is disturbed. By analyzing the timing of kinetochore separation, we find that this anaphase delay requires the Bub3, Mad3, and Bub1 but not the Mad1 or Mad2 spindle assembly checkpoint proteins. In agreement with this, we find that Bub1 remains associated with kinetochores when spindles are mispositioned. These data indicate that, in fission yeast, astral microtubule contact with the medial cell cortex is monitored by a subset of spindle assembly checkpoint proteins. We propose that this checkpoint ensures spindles are properly oriented before anaphase takes place.

Online version of this article contains supporting material. Online version is available at www.molbiolcell.org.

These authors contributed equally to this work.

Present address: Le Laboratoire de Biologie Cellulaire et Moléculaire du Contrôle de la Prolifáeration-Centre National de la Recherche Scientifique UMR5088, Institut d'Exploration Fonctionelle des Génomes (IFR109), Université Paul Sabatier, 118 route de Narbonne, 31062 Toulouse, France.

^|| Corresponding author. E-mail address: jmillar{at}nimr.mrc.ac.uk.

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