Auxin at the Shoot Apical Meristem

Plants continuously generate new tissues and organs through the activity of populations of undifferentiated stem cells, called meristems. Here, we discuss the so-called shoot apical meristem (SAM), which generates all the aerial parts of the plant. It has been known for many years that auxin plays a central role in the functioning of this meristem. Auxin is not homogeneously distributed at the SAM and it is thought that this distribution is interpreted in terms of differential gene expression and patterned growth. In this context, auxin transporters of the PIN and AUX families, creating auxin maxima and minima, are crucial regulators. However, auxin transport is not the only factor involved. Auxin biosynthesis genes also show specific, patterned activities, and local auxin synthesis appears to be essential for meristem function as well. In addition, auxin perception and signal transduction defining the competence of cells to react to auxin, add further complexity to the issue. To unravel this intricate signaling network at the SAM, systems biology approaches, involving not only molecular genetics but also live imaging and computational modeling, have become increasingly important.Plants continuously generate new tissues and organs through the activity of populations of undifferentiated stem cells, called meristems. Because meristems can modulate their activity, they provide the developmental flexibility that allows plants to adapt their development in reaction to the environment (reviews: Lyndon 1998; Traas and Doonan 2001; Aida and Tasaka 2006; Sablowski 2007).Distinct meristems exist. Apical meristems, positioned at the tip of the shoots and roots, initiate aerial and underground organs, respectively. Along the stems and roots, more diffuse secondary meristems are responsible for secondary thickening of these structures.The plant hormone auxin plays an instrumental role in meristem biology and we discuss here its role in a particular meristem, the shoot apical meristem (SAM) that generates all the aerial organs including the floral meristems (Fig. 1A–C). In this context, we limit ourselves to the meristems in angiosperm that have been studied in most detail.Open in a separate windowFigure 1.The shoot apical meristem of Arabidopsis thaliana. (A) Aerial part of a wild-type plant of the Columbia ecotype (Col-0). The SAM is responsible for the production of rosette leaves and, after floral transition, for the production of the stem, cauline leaves, lateral meristems, and flowers of the inflorescence. (B) Details of the tip of the inflorescence, showing the highly organized positioning of flowers around the main axis (a spiral). (C) A dissected inflorescence meristem. Older flowers have been removed to expose the meristem surrounded by young floral buds. (D) Longitudinal section of an inflorescence meristem showing the layered organization (L1, L2, and L3 cell layers). L1 and L2 are also called the tunica and L3 to the corpus. The functional zones are also represented. At the meristem summit the central zone (CZ) contains the stem cells, whereas primordia are initiated in the peripheral zone (PZ). The rib zone (RZ) produces the internal part of the stem.