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Vol. 11, Issue 11, 3949-3961, November 2000
and
*Department of Biology, University of North Carolina at Chapel
Hill, Chapel Hill, North Carolina 27599-3280; In the budding yeast Saccharomyces cerevisiae,
movement of the mitotic spindle to a predetermined cleavage plane at
the bud neck is essential for partitioning chromosomes into the mother and daughter cells. Astral microtubule dynamics are critical to the
mechanism that ensures nuclear migration to the bud neck. The nucleus
moves in the opposite direction of astral microtubule growth in the
mother cell, apparently being "pushed" by microtubule contacts at
the cortex. In contrast, microtubules growing toward the neck and
within the bud promote nuclear movement in the same direction of
microtubule growth, thus "pulling" the nucleus toward the bud neck.
Failure of "pulling" is evident in cells lacking Bud6p, Bni1p,
Kar9p, or the kinesin homolog, Kip3p. As a consequence, there is a loss
of asymmetry in spindle pole body segregation into the bud. The
cytoplasmic motor protein, dynein, is not required for nuclear movement
to the neck; rather, it has been postulated to contribute to spindle
elongation through the neck. In the absence of KAR9,
dynein-dependent spindle oscillations are evident before anaphase
onset, as are postanaphase dynein-dependent pulling forces that exceed
the velocity of wild-type spindle elongation threefold. In addition,
dynein-mediated forces on astral microtubules are sufficient to
segregate a 2N chromosome set through the neck in the absence of
spindle elongation, but cytoplasmic kinesins are not. These
observations support a model in which spindle polarity determinants
(BUD6, BNI1, KAR9) and
cytoplasmic kinesin (KIP3) provide directional cues for
spindle orientation to the bud while restraining the spindle to the
neck. Cytoplasmic dynein is attenuated by these spindle polarity
determinants and kinesin until anaphase onset, when dynein directs
spindle elongation to distal points in the mother and bud.
Department of
Pharmacology and Cancer Biology, Duke University Medical Center,
Durham, North Carolina 27709
Online version of this article contains video material.
Corresponding author. E-mail address:
kerry_bloom{at}unc.edu.
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