Formation of aperture sites on the pollen surface as a model for development of distinct cellular domains

Pollen grains are covered by the complex extracellular structure, called exine, which in most species is deposited on the pollen surface non-uniformly. Certain surface areas receive fewer exine deposits and develop into regions whose structure and morphology differ significantly from the rest of pollen wall. These regions are known as pollen apertures. Across species, pollen apertures can vary in their numbers, positions, and morphology, generating highly diverse patterns. The process of aperture formation involves establishment of cell polarity, formation of distinct plasma membrane domains, and deposition of extracellular materials at precise positions. Thus, pollen apertures present an excellent model for studying the development of cellular domains and formation of patterns at the single-cell level. Until very recently, the molecular mechanisms underlying the specification and formation of aperture sites were completely unknown. Here, we review recent advances in understanding of the molecular processes involved in pollen aperture formation, focusing on the molecular players identified through genetic approaches in the model plant Arabidopsis. We discuss a potential working model that describes the process of aperture formation, including specification of domains, creation of their defining features, and protection of these regions from exine deposition.