TheHandlebarsgene is required withPhantasticafor dorsoventral asymmetry of organs and for stem cell activity inAntirrhinum

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Abstract
In angiosperms, individual lateral organs and whole flowers may develop asymmetrically along their dorsoventral axes. Dorsoventral asymmetry of Antirrhinum leaves requires activity of the Phantastica gene and other factors acting redundantly with it. We describe the effects of a mutation in the Handlebars gene, identified as an enhancer of the phantastica mutant phenotype. Genetic analysis suggests that Handlebars functions redundantly with Phantastica to promote dorsal fate in lateral organs and to maintain activity of stem cells within shoot apical meristems. Handlebars appears dispensable in vegetative development but is needed for asymmetry of petals along the dorsoventral axis of the flower as a whole. This suggests that common mechanisms may control dorsoventral asymmetry in lateral organ primordia and in floral meristems.