Pattern Formation and Complexity in Single Cells.

In the context of animal or plant development, we tend to think of cells as small, simple, building blocks, such that complex patterns or shapes can only be constructed from large numbers of cells, with cells in different parts of the organism taking on different fates. However, cells themselves are far from simple, and often take on complex shapes with a remarkable degree of intracellular patterning. How do these patterns arise? As in embryogenesis, the development of structure inside a cell can be broken down into a number of basic processes. For each part of the cell, morphogenetic processes create internal structures such as organelles, which might correspond to organs at the level of a whole organism. Given that mechanisms exist to generate parts, patterning processes are required to ensure that the parts are distributed in the correct arrangement relative to the rest of the cell. Such patterning processes make reference to global polarity axes, requiring mechanisms for axiation which, in turn, require processes to break symmetry. These fundamental processes of symmetry breaking, axiation, patterning, and morphogenesis have been extensively studied in developmental biology but less so at the subcellular level. This review will focus on developmental processes that give eukaryotic cells their complex structures, with a focus on cytoskeletal organization in free-living cells, ciliates in particular, in which these processes are most readily apparent.