Spindle orientation processes in epithelial growth and organisation.

This review focuses on the role of orientated cell division (OCD) in two aspects of epithelial growth, namely layer formation and growth in the epithelial plane. Epithelial stratification is invariably associated with fate asymmetric cell divisions. We discuss this through the example of epidermal stratification where cell division plane regulation facilitates concomitant thickening and cell differentiation. Embryonic neuroepithelia are considered as a special case of epithelial stratification. We highlight early ectodermal layer specification, which sets the epidermal versus neuronal fates, as well as later neurogenesis in vertebrates and mammals. We also discuss the heart epicardium as an example of coordinating OCDs with delamination and subsequent differentiation. Epithelial planar growth is examined both in the context of uniform growth, such as in Xenopus epiboly, the Drosophila wing disc and the mammalian intestinal crypt as well as in anisotropic growth, or elongation, such as Drosophila and vertebrate axial elongation and the mouse palate. Coupling between growth perpendicular to and within epithelial planes is recognised, but so are exceptions, as is the often passive role of spindle orientation sometimes hitherto considered to be an active driver of directional growth.