Traveling Waves Enhance Transport during Cytoplasmic Streaming in Drosophila Oogenesis

Cytoplasmic streaming is believed crucial to large developing cells, such as oocytes, where flow provides a much faster route to transport and mixing of cellular components than molecular diffusion. Molecular motors carrying cargoes on cytoskeletal elements are known to drive streaming in a variety of biological systems and, in Drosophila oocytes, cell-spanning vortical flows are tied to kinesin motors moving on dense beds of microtubules. Recent theories and simulations suggest such streaming self-organizes through interacting microtubules, motors, and flow. Whether these theoretically derived flows are adequate to the tasks of transport and mixing has been unclear. Here we report on new observations of coherent waves traveling persistently through microtubule beds during streaming, and investigate their impact on fluid transport and mixing. Leveraging recent advances in simulating the complex fluid-structure problems of cellular environments, we probe a biophysical model set in an oocytal geometry. Our simulations identify previously unknown waving states that show concordance with our experimental observation and the capacity for functional transport and mixing.