The invariant cleavage pattern displayed by ascidian embryos depends on spindle positioning along the cell's longest axis in the apical plane and relies on asynchronous cell divisions

The position of cells within an embryo early in development determines what type of cells they will become in the fully formed embryo. The embryos of ascidians, commonly known as sea squirts, are ideal for studying what influences cell positioning. These embryos consist of a small number of cells that divide according to an “invariant cleavage pattern”, which means that the positioning and timing of the cell divisions is identical between different individuals of the same species. The pattern of cell division is also largely the same across different ascidian species, which raises questions about how it is controlled. When a cell divides, a structure called the spindle forms inside it to distribute copies of the cell’s genetic material between the new cells. The orientation of the spindle determines the direction in which the cell will divide. Now, by combining 3D imaging of living ascidian embryos with computational modeling, Dumollard et al. show that the spindles in every equally dividing cell tend to all align in the long length of the cell’s “apical” surface. Such alignment allows the cells to be on the outside of the embryo and implements the ascidian invariant cleavage pattern. The cells in the embryo do not all divide at the same time. Indeed, the shape of the cells (and especially their apical surface) depends on two layers of cells in the embryo not dividing at the same time; instead, periods of cell division alternate between the layers. A network of genes in the embryo regulates the timing of these cell divisions and makes it possible for the cells to divide according to an invariant cleavage pattern. However, this network of genes is not the only control mechanism that shapes the early embryo. A structure found in egg cells (and hence produced by the embryo’s mother) causes cells at the rear of the embryo to divide unequally, and this influences the shape of all the cells in the embryo. Thus it appears that maternal mechanisms work alongside the embryo’s gene network to shape the early embryo. The next step will be to determine how physical forces – for example, from the cells pressing against each other – influence the position of the embryo’s cells. How do gene networks relay the biomechanical properties of the embryo to help it take shape?