Polarity and axis formation in the Drosophila female germ line

By the time a Drosophila egg is laid, both major body axes have already been defined and it contains all the nutrients needed to develop into a free-living larva in 24 h. By contrast, it takes almost a week to make an egg from a female germline stem cell, during the complex process of oogenesis. This review will discuss key symmetry-breaking steps in Drosophila oogenesis that lead to the polarisation of both body axes: the asymmetric divisions of the germline stem cells; the selection of the oocyte from the 16-cell germline cyst; the positioning of the oocyte at the posterior of the cyst; Gurken signalling from the oocyte to polarise the anteriorposterior axis of the somatic follicle cell epithelium around the developing germline cyst; the signalling back from the posterior follicle cells to polarise the anterior-posterior axis of the oocyte; and the migration of the oocyte nucleus that specifies the dorsal-ventral axis. Since each event creates the preconditions for the next, I will focus on the mechanisms that drive these symmetry-breaking steps, how they are linked and the outstanding questions that remain to be answered.