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Vol. 16, Issue 8, 3666-3677, August 2005

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Spindle Checkpoint Signaling Requires the Mis6 Kinetochore Subcomplex, Which Interacts with Mad2 and Mitotic Spindles

Shigeaki Saitoh ^*, Kojiro Ishii ^*, Yasuyo Kobayashi ^*, and Kohta Takahashi ^* 

^* Division of Cell Biology, Institute of Life Science, Kurume University, Fukuoka 839-0864, Japan; Time's Arrow and Biosignaling, Precursory Research for Embryonic Science and Technology, Japanese Science and Technology, Saitama 332-0012, Japan

Submitted January 7, 2005; Revised April 28, 2005; Accepted May 24, 2005
Monitoring Editor: Ted Salmon

The spindle checkpoint coordinates cell cycle progression and chromosome segregation by inhibiting anaphase promoting complex/cyclosome until all kinetochores interact with the spindle properly. During early mitosis, the spindle checkpoint proteins, such as Mad2 and Bub1, accumulate at kinetochores that do not associate with the spindle. Here, we assess the requirement of various kinetochore components for the accumulation of Mad2 and Bub1 on the kinetochore in fission yeast and show that the necessity of the Mis6-complex and the Nuf2-complex is an evolutionarily conserved feature in the loading of Mad2 onto the kinetochore. Furthermore, we demonstrated that Nuf2 is required for maintaining the Mis6-complex on the kinetochore during mitosis. The Mis6-complex physically interacts with Mad2 under the condition that the Mad2-dependent checkpoint is activated. Ectopically expressed N-terminal fragments of Mis6 localize along the mitotic spindle, highlighting the potential binding ability of Mis6 not only to the centromeric chromatin but also to the spindle microtubules. We propose that the Mis6-complex, in collaboration with the Nuf2-complex, monitors the spindle–kinetochore attachment state and acts as a platform for Mad2 to accumulate at unattached kinetochores.

Abbreviations used: APC/C, anaphase promoting complex/cyclosome; CBZ, carbentazim; MT, microtubule; SPB, spindle pole body.

The online version of this article contains supplemental material at MBC Online (http://www.molbiolcell.org).

Address correspondence to: Kohta Takahashi (takahash{at}lsi.kurume-u.ac.jp).

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