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A more recent version of this article appeared on August 1, 2005
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Submitted on April 3, 2005
Revised on May 20, 2005
Accepted on May 23, 2005
*Department of Biomedical Engineering, University of Minnesota, Minneapolis, MN 55455; Department of Biology, University of North Carolina at Chapel Hill, Chapel Hill, NC 27599; Lab of Receptor Biology and Gene Expression, National Cancer Institute, Bethesda MD 20892
Monitoring Editor: Orna Cohen-Fix
During metaphase in budding yeast mitosis, sister kinetochores are tethered to opposite poles and separated, stretching their intervening chromatin, by singly attached kinetochore microtubules (kMTs). Kinetochore movements are coupled to single microtubule plus-end polymerization/depolymerization at kinetochore attachment sites. Here we use computer modeling to test possible mechanisms controlling chromosome alignment during yeast metaphase by simulating experiments that determine: 1) mean positions of kinetochore Cse4-GFP, 2) the extent of oscillation of kinetochores during metaphase as measured by fluorescence recovery after photobleaching (FRAP) of kinetochore Cse4-GFP, 3) the dynamics of kMTs as measured by FRAP of GFP-tubulin, and 4) mean positions of unreplicated-chromosome kinetochores that lack pulling forces from a sister kinetochore. We rule out a number of possible models and find the best fit between theory and experiment when it is assumed that kinetochores sense both a spatial gradient that suppresses kMT catastrophe near the poles, and attachment site tension that promotes kMT rescue at higher amounts of chromatin stretch.
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