Hormonal regulation of cereal endosperm development with a focus on rice (Oryza sativa)

The endosperm of cereal grain forms the staple diet for most of the world's population, and feeds much of their stock. Grain size and quality are determined largely by events taking place during coenocytic nuclear division, endosperm cellularisation and cell differentiation, and the production of storage molecules. Thus, understanding the complex signalling processes occurring at each of these steps is essential for maintaining and improving our food supply. Here, we critically review evidence for the effects of phytohormones on grain size, as well as hormone homeostasis, signalling and crosstalk. We focus on rice endosperm due to the importance of rice as a food crop and a model grass, as well as its relative neglect in recent reviews; however, data from other cereals are also discussed due to strong evidence for conserved signalling networks operating during grain development. Discussion is restricted to auxin, cytokinin, ethylene, abscisic acid and gibberellin. Our review highlights the need for accurate hormone determinations combined with information on gene expression. We present evidence for separate, localised signalling roles for auxin at different stages of grain development and highlight key research questions for other hormones where much less data are available.