The genesis of retinal architecture: an emerging role for mechanical

Patterns in nature have always fascinated human beings. They convey the idea of order, organization and optimization, and, to the enquiring mind, the alluring promise that understanding their building rules may uncover the forces that shaped them. In the retina, two patterns are outstanding: the stacking of cells in layers and, within the layers, the prevalent arrangement of neurons of the same type in orderly arrays, often referred to as mosaics for the crystalline-like order that some can display. Layers and mosaics have been essential keys to our present understanding of retinal circuital organization and function. Now, they may also be a precious guide in our exploration of how the retina is built. Here, we will review studies addressing the mechanisms controlling the formation of retinal mosaics and layers, illustrating common themes and unsolved problems. Among the intricacies of the building process, a world of physical forces is making its appearance. Cells are extremely complex to model as ''physical entities'', and many aspects of cell mechanotransduction are still obscure. Yet, recent experiments, focusing on the mechanical aspects of growth and differentiation, suggest that adopting this viewpoint will open new ways of understanding retinal formation and novel possibilities to approach retinal pathologies and repair.