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A more recent version of this article appeared on December 1, 2002
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Submitted on June 17, 2002
Revised on August 20, 2002
Accepted on August 23, 2002
1 European Molecular Biology Laboratory, Meyerhofstrasse 1, 69117 Heidelberg, Germany
2 European Molecular Biology Laboratory, Meyerhofstrasse 1, 69117 Heidelberg, Germany (present address: Max Planck Institute of Molecular Cell Biology and Genetics, 01307 Dresden, Germany)
* Corresponding author. E-mail address: mattaj{at}embl-heidelberg.de.
The small GTPase Ran has been found to play pivotal roles in several aspects of cell function. We have investigated the role of the Ran GTPase cycle in spindle formation and nuclear envelope assembly in dividing Caenorhabditis elegans embryos in real time. We found that Ran and its co-factors RanBP2, RanGAP and RCC1 are all essential for reformation of the nuclear envelope after cell division. Reducing the expression of any of these components of the Ran GTPase cycle by RNAi leads to strong extranuclear clustering of integral nuclear envelope proteins and nucleoporins. Ran, RanBP2, and RanGAP are also required for building a mitotic spindle, whereas astral microtubules are normal in the absence of these proteins. RCC1(RNAi) embryos have similar abnormalities in the initial phase of spindle formation but eventually recover to form a bipolar spindle. Irregular chromatin structures and chromatin bridges due to spindle failure were frequently observed in embryos where the Ran cycle was perturbed. In addition, connection between the centrosomes and the male pronucleus, and thus centrosome positioning, depend upon the Ran cycle components. Finally, we have demonstrated that both IMA-2 and IMB-1, the homologues of vertebrate importin 
and ß, are essential for both spindle assembly and nuclear formation in early embryos.
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