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MBC in Press, published online ahead of print August 3, 2005
Mol. Biol. Cell 10.1091/mbc.E05-02-0154

A more recent version of this article appeared on October 1, 2005
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Submitted on February 23, 2005
Revised on June 10, 2005
Accepted on July 21, 2005

Early Spindle Assembly in Drosophila Embryos: Role of a Force Balance Involving Cytoskeletal Dynamics and Nuclear Mechanics

E. N. Cytrynbaum,*{dagger}{ddagger} P. Sommi,*{ddagger} I. Brust-Mascher,* J. M. Scholey,*{sect}|| and A. Mogilner*||¶

*Laboratory of Cell and Computational Biology, Center for Genetics and Development, University of California-Davis, Davis, CA 95616; {dagger}Department of Mathematics, University of British Columbia, Vancouver, British Columbia V6T 1Z2, Canada; {sect}Section of Molecular and Cell Biology and Department of Mathematics, University of California-Davis, Davis, CA 95616

Monitoring Editor: Ted Salmon

Mitotic spindle morphogenesis depends upon the action of microtubules (MTs), motors and the cell cortex. Previously we proposed that cortical- and MT-based motors acting alone can coordinate early spindle assembly in Drosophila embryos. Here we tested this model using microscopy of living embryos to analyze spindle pole separation, cortical reorganization and nuclear dynamics in interphase-prophase of cycles 11-13. We observe that actin caps remain flat as they expand, and that furrows do not ingress. As centrosomes separate they follow a linear trajectory, maintaining a constant pole-to-furrow distance while the nucleus progressively deforms along the elongating pole-pole axis. These observations are incorporated into a model in which outward forces generated by zones of active cortical dynein are balanced by inward forces produced by nuclear elasticity and, during cycle 13, by Ncd, which localizes to interpolar MTs. Thus, the force-balance driving early spindle morphogenesis depends upon MT-based motors acting in concert with the cortex and nucleus.

{ddagger}These authors contributed equally to this work.

||These authors are codirectors of this project.

Address correspondence to: A. Mogilner (mogilner{at}math.ucdavis.edu)




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