Cereal Grass Pulvini: Agronomically Significant Models for Studying Gravitropism Signaling and Tissue Polarity

Cereal grass pulvini have emerged as model systems that are not only valuable for the study of gravitropism, but are also of agricultural and economic significance. The pulvini are regions of tissue that are apical to each node and collectively return a reoriented stem to a more vertical position. They have proven to be useful for the study of gravisensing and response and are also providing clues about the establishment of polarity across tissues. This review will first highlight the agronomic significance of these stem regions and their benefits for use as model systems and provide a brief historical overview. A detailed discussion of the literature focusing on cell signaling and early changes in gene expression will follow, culminating in a temporal framework outlining events in the signaling and early growth phases of gravitropism in this tissue. Changes in cell wall composition and gene expression that occur well into the growth phase will be touched upon briefly. Finally, some ongoing research involving both maize and wheat pulvini will be introduced along with prospects for future investigations.