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We thank Mitsuhiro Aida for sharing cuc1 and cuc2 seeds prior to publication, Ki-Young Suh for help with the enhancer screening, Michael Frohlich for discussions about filamentous structures, and Pat Coen and Jean Edens of the Caltech Electron Microscope Facility for assistance with SEM and TEM. Plants doubly mutant for an ffo gene and for ufo, lfy, or clv3 reveal that the FFO genes play roles related to those of UFO and LFY in floral meristem initiation and that FFO2 and FFO3 may act to control cell proliferation late in inflorescence development. Plants doubly mutant for two ffo alleles display additive phenotypes, indicating that the FFO genes may act in separate pathways.

FFO2 acts at early floral stages to regulate floral organ number and positioning and to control organ separation within and between whorls. FFO1 and FFO3 have specific functions in cauline leaf/stem separation and in first- and third-whorl floral organ separation, with FFO3 likely acting to establish and FFO1 to maintain floral organ boundaries. ffo mutants have specific defects in floral organ separation and/or positioning thus, the FFO genes identify components of a boundary formation mechanism(s)acting between developing floral organ primordia. The mutants fall into three complementation groups, which we have termed FUSED FLORAL ORGANS (FFO) loci. In a screen to identify novel genes required for early Arabidopsis flower development, we isolated four independent mutations that enhance the Ufo phenotype toward the production of filamentous structures in place of flowers.