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First published online August 2, 2005
doi: 10.1242/10.1242/dev.01956
1 Department of Biology, Indiana University, Bloomington, 1001 East 3rd Street,
IN 47405, USA
2 Program in Cell Dynamics, University of Massachusetts, 55 Lake Avenue, North
Worcester, MA 01655, USA
* Author for correspondence (e-mail: bsaxton{at}bio.indiana.edu)
Accepted 22 June 2005
Mass movements of cytoplasm, known as cytoplasmic streaming, occur in some large eukaryotic cells. In Drosophila oocytes there are two forms of microtubule-based streaming. Slow, poorly ordered streaming occurs during stages 8-10A, while pattern formation determinants such as oskar mRNA are being localized and anchored at specific sites on the cortex. Then fast well-ordered streaming begins during stage 10B, just before nurse cell cytoplasm is dumped into the oocyte. We report that the plus-end-directed microtubule motor kinesin-1 is required for all streaming and is constitutively capable of driving fast streaming. Khc mutations that reduce the velocity of kinesin-1 transport in vitro blocked streaming yet still supported posterior localization of oskar mRNA, suggesting that streaming is not essential for the oskar localization mechanism. Inhibitory antibodies indicated that the minus-end-directed motor dynein is required to prevent premature fast streaming, suggesting that slow streaming is the product of a novel dynein-kinesin competition. As F-actin and some associated proteins are also required to prevent premature fast streaming, our observations support a model in which the actin cytoskeleton triggers the shift from slow to fast streaming by inhibiting dynein. This allows a cooperative self-amplifying loop of plus-end-directed organelle motion and parallel microtubule orientation that drives vigorous streaming currents and thorough mixing of oocyte and nurse-cell cytoplasm.
Key words: Drosophila, Oocyte, Kinesin-1, Dynein, Streaming, Microtubule, Actin, Oskar
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