Perspectives on leaf dorsoventral polarity

Leaves occur in a vast array of shapes and sizes, with complex diversity contributing to optimization of the principal function of photosynthesis. The program of development from a self-renewing stem cell population to a mature leaf has been of interest to biologists for years. Many genes involved in this process have been identified, particularly in the model eudicot Arabidopsis, so that now we have a greater understanding of mechanisms of stem cell maintenance, cell differentiation and organogenesis. One aspect of leaf development that is of particular interest is the establishment of dorsoventral polarity: the distinct adaxial (upper) and abaxial (lower) sides of the leaf. Early studies postulated conceptual models of how establishment of polarity leads to the development of planar leaves. Studies over the past decade have defined genetic details of this model, and uncovered diverse mechanisms of gene regulation that facilitate development of leaf dorsoventral polarity, including transcriptional regulation, chromatin modification, DNA modification, regulation by short RNAs and translational and post-translational regulation. This review will discuss these regulatory mechanisms in the context of leaf dorsoventrality, and will conclude with unresolved questions and areas of future research.