A naturally occurring frameshift mutation in the UNUSUAL FLORAL ORGANS gene associated with the marimo floral phenotype in gerbera
Background The capitulum of Asteraceae is a highly specialized inflorescence whose formation requires the coordinated regulation of multiple developmental processes, including floral organ identity and floral meristem determinacy. The LEAFY (LFY)–UNUSUAL FLORAL ORGANS (UFO) regulatory module is known to play an important role in flower development; however, naturally occurring mutations affecting this pathway have not been genetically characterized in gerbera (Gerbera hybrida). Results In this study, we characterized a novel gerbera mutant identified during a commercial crossing program and named it marimo based on its green, spherical capitulum. Morphological observations revealed the repeated formation of secondary and tertiary floret-like organs within primary floret-like organs. Scanning electron microscopy showed that the epidermal structure of the green organs in marimo was similar to that of wild-type involucral bracts. RNA sequencing identified numerous differentially expressed genes between marimo and the wild type, and network and Gene Ontology analyses highlighted gene groups associated with flower development, reproductive organ differentiation, and tissue structure formation. RNA-seq analysis showed increased expression of LFY and reduced expression of GGLO1, a PISTILLATA/GLOBOSA-like B-class MADS-box gene, in the marimo mutant. RT-qPCR analysis of a segregating population further confirmed reduced GGLO1 expression in marimo-type individuals. In addition, a single-nucleotide deletion was identified in the coding region of UFO. This deletion was predicted to cause a frameshift and a premature stop codon. In selfed progeny of No. 251, the UFO genotype was fully associated with capitulum phenotype, and only individuals homozygous for the mutant allele exhibited the marimo phenotype. Conclusions These results indicate that the naturally occurring frameshift mutation in UFO is the strongest candidate variant underlying the marimo phenotype. RNA-seq analysis showed increased LFY expression and markedly reduced GGLO1 expression in the marimo mutant. Reduced activity of the LFY–UFO regulatory module may therefore have altered the expression of GGLO1 and other floral organ development-related genes despite the continued expression of LFY. These changes may have affected both floral organ identity and floral meristem determinacy, resulting in the formation of green involucral bract-like organs and the repeated production of floret-like organs. The marimo mutant provides a useful genetic resource for investigating capitulum development in Asteraceae and may also serve as breeding material for introducing novel ornamental traits into gerbera.