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Published online 18 October 2004. doi:10.1083/jcb.200406008
The Rockefeller University Press, 0021-9525 $8.00
JCB, Volume 167, Number 2, 245-256
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Article

The forces that position a mitotic spindle asymmetrically are tethered until after the time of spindle assembly



Jean-Claude Labbé, Erin K. McCarthy, and Bob Goldstein

Department of Biology, The University of North Carolina at Chapel Hill, Chapel Hill, NC 27599

Correspondence to J.-C. Labbé: jc.labbe{at}umontreal.ca

Regulation of the mitotic spindle's position is important for cells to divide asymmetrically. Here, we use Caenorhabditis elegans embryos to provide the first analysis of the temporal regulation of forces that asymmetrically position a mitotic spindle. We find that asymmetric pulling forces, regulated by cortical PAR proteins, begin to act as early as prophase and prometaphase, even before the spindle forms and shifts to a posterior position. The spindle does not shift asymmetrically during these early phases due to a tethering force, mediated by astral microtubules that reach the anterior cell cortex. We show that this tether is normally released after spindle assembly and independently of anaphase entry. Monitoring microtubule dynamics by photobleaching segments of microtubules during anaphase revealed that spindle microtubules do not undergo significant poleward flux in C. elegans. Together with the known absence of anaphase A, these data suggest that the major forces contributing to chromosome separation during anaphase originate outside the spindle. We propose that the forces positioning the mitotic spindle asymmetrically are tethered until after the time of spindle assembly and that these same forces are used later to drive chromosome segregation at anaphase.

J.-C. Labbé's present address is Swiss Federal Institute of Technology, Institute of Biochemistry, ETH Hönggerberg HPM G16, 8093 Zürich, Switzerland.

Abbreviations used in this paper: c-LßL, clasto-lactacystin ß-lactone; EL, embryonic length; OICD, optically induced centrosome disruption.


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