Lateral plant organs, particularly leaves, initiate at the flanks of the shoot apical meristem (SAM) following auxin maxima signals; however, little is known about the underlying mechanisms. Here, we show that tomato leafless (lfs) mutants fail to produce cotyledons and leaves and grow a naked pin while maintaining an active SAM. A similar phenotype was observed among pin-like shoots induced by polar auxin transport inhibitors such as 2,3,5-triiodobenzoic acid (TIBA). Both types of pin-like shoots showed reduced expression of primordia markers as well as abnormal auxin distribution, as evidenced by expression of the auxin reporters pPIN1: PIN1:GFP and DR5:YFP. Upon auxin microapplication, both lfs meristems and TIBA-pin apices activated DR5:YFP expression with similar kinetics; however, only lfs plants failed to concurrently initiate leaf primordia. We found that LFS encodes the single tomato ortholog of Arabidopsis DORNRONSCHEN (DRN) and DRN-like (DRNL) genes and is transiently expressed at incipient and young primordia, overlapping with auxin response maxima. LFS is rapidly induced by auxin application, implying feed-forward activity between LFS and auxin signals. However, driving LFS at auxin response maxima sites using the DR5 promoter fails to fully rescue lfs plants, suggesting that additional, auxin-independent regulation is needed. Indeed, extended GCC-box elements upstream of LFS drove primordia-specific expression in a LFS-dependent but auxin-independent manner. We thus suggest that LFS transiently acts at the site of primordia initiation, where it provides a specific context to auxin response maxima culminating in leaf primordia initiation.L ateral aerial plant organs form at the flanks of the shoot apical meristem (SAM) in a species-specific order termed phyllotaxis. Phyllotactic patterns tightly correlate with local maxima of the phytohormone auxin at the SAM periphery (1-3). These maxima are achieved by active transport (4). At these sites, auxin is required for leaf initiation, and when polar auxin transport is chemically impaired by polar auxin transport inhibitors (PATI), leaves fail to form (5-7), resulting in a pin-like shoot. When auxin is externally applied onto the meristem of these pins or to the meristem in a wild-type (WT) background, leaf primordia are initiated (5). Genetic analyses have also demonstrated that auxin response maxima, dictated by the activity of the auxin efflux carrier PINFORMED1 (PIN1), can be detected before, and are required for, leaflet initiation in compound leaves (6). However, similar evidence for leaf initiation is lacking (7).Like other classical hormones, such as cytokinin (CK), florigen, and gibberellin, auxin has many functions, which are largely dictated by its levels and which vary within and among plants. In young leaf primordia, auxin triggers primordia bulging at P 0 by stimulating local cell divisions. At P 1 and P 2 , auxin elicits provasculature development (8). Thus, in the same group of cells, auxin cues are differentially translated to cell divisi...