Master transcription factors reprogram cell fate in multicellular eukaryotes. Pioneer transcription factors have prominent roles in this process because of their ability to contact their cognate binding motifs in closed chromatin. Reprogramming is pervasive in plants, whose development is plastic and tuned by the environment, yet no bonafide pioneer transcription factor has -been identified in this kingdom. Here we show that the master transcription factor LEAFY (LFY), which promotes floral fate through upregulation of the floral commitment factor APETALA1 (AP1), is a pioneer transcription factor. In vitro, LFY binds in a sequence-specific manner and with high affinity to the endogenous AP1 target locus DNA assembled into a nucleosome. In vivo, LFY associates with nucleosome occupied binding sites at the majority of its target loci, including AP1, where it co-occupies DNA with histones. Moreover, the LFY DNA contact helix shares defining properties with those of strong nucleosome binding pioneer factors. At the AP1 locus, LFY unlocks chromatin locally by displacing the H1 linker histone and by recruiting SWI/SNF chromatin remodelers, but broad changes in chromatin accessibility occur later and require activity of additional, non-pioneer transcription, factors. Our study provides a mechanistic framework for patterning of inflorescence architecture and uncovers striking similarities between plant and animal pioneer transcription factors. Further analyses aimed at elucidating the defining characteristics of pioneer transcription factors will allow harnessing these for enhanced cell fate reprogramming.Chromatin prevents expression of inappropriate or detrimental gene expression programs, allowing formation of distinct cell types from the same genome 1. The basic unit of chromatin is the nucleosome comprised of 150 base-pairs of DNA wrapped nearly two turns around the histone octamer 2. Chromatin is further compacted by nucleosome/nucleosome interactions and by the linker histone H1, which associates with the dyad (midpoint) of the nucleosome 3. During cell fate reprogramming in eukaryotes, master transcription factors silence and activate new gene expression programs in the context of chromatin 4-6. While it is easy to imagine how master transcription factors bind to active genes in open chromatin to trigger silencing, it is difficult to envision how these sequence-specific binding proteins can access their cognate motifs in silent chromatin to activate gene expression. This is because nucleosomes are refractory for most transcription factor binding 7-10 11. However, a special class of transcription factors, termed pioneer transcription factors, can access their cognate binding motifs in the nucleosome 1,4,12-17. These pioneer transcription factors play important roles in cell fate reprogramming. For example, the mammalian pioneer transcription factor FoxA reprograms fibroblast to hepatocytes 16,18-20, while the Oct4, Klf4 and Sox2 pioneer transcription factors reprogram fibroblasts to induced pluripotent stem ...