Leaves originate from the shoot apical meristem, a small mound of undifferentiated tissue at the tip of the stem. Leaf formation begins with the selection of a group of founder cells in the so-called peripheral zone at the flank of the meristem, followed by the initiation of local growth and finally morphogenesis of the resulting bulge into a differentiated leaf. Whereas the mechanisms controlling the switch between meristem propagation and leaf initiation are being identified by genetic and molecular analyses, the radial positioning of leaves, known as phyllotaxis, remains poorly understood. Hormones, especially auxin and gibberellin, are known to influence phyllotaxis, but their specific role in the determination of organ position is not clear. We show that inhibition of polar auxin transport blocks leaf formation at the vegetative tomato meristem, resulting in pinlike naked stems with an intact meristem at the tip. Microapplication of the natural auxin indole-3-acetic acid (IAA) to the apex of such pins restores leaf formation. Similarly, exogenous IAA induces flower formation on Arabidopsis pin-formed1-1 inflorescence apices, which are blocked in flower formation because of a mutation in a putative auxin transport protein. Our results show that auxin is required for and sufficient to induce organogenesis both in the vegetative tomato meristem and in the Arabidopsis inflorescence meristem. In this study, organogenesis always strictly coincided with the site of IAA application in the radial dimension, whereas in the apical-basal dimension, organ formation always occurred at a fixed distance from the summit of the meristem. We propose that auxin determines the radial position and the size of lateral organs but not the apical-basal position or the identity of the induced structures.
INTRODUCTIONLeaf primordia develop in regular patterns from the shoot apical meristem. In the majority of flowering plants, the leaves are arranged in spirals, with the divergence angle between successive leaves approaching the Fibonacci angle of 137.5 Њ , the so-called golden ratio (Steeves and Sussex, 1989;Lyndon, 1990Lyndon, , 1998Jean, 1994). Molecular and genetic analyses have recently identified various genes that regulate meristem development and leaf formation. Some of these genes encode transcription factors of the homeobox class (Vollbrecht et al., 1991;Long et al., 1996;Kerstetter et al., 1997; Taylor, 1997;Mayer et al., 1998), whereas others, such as clavata1 , clavata3, and zwille, appear to be involved in cell-to-cell signaling Moussian et al., 1998;Fletcher et al., 1999; Trotochaud et al., 1999). Current models postulate that these genes control the balance between meristem self-propagation and the production of organogenic tissue.Whereas genetic analyses provide us with an ever more detailed description of meristem maintenance and propagation, the molecular nature of the mechanisms that trigger leaf initiation and ensure that leaves form at the proper angles relative to each other remains to be established. In many m...