The Gibberellin perception system evolved to regulate a pre-existing GAMYB-mediated system during land plant evolution

Gibberellin (GA) controls pollen development in flowering plants via the GAMYB transcription factor. Here we show that GAMYB is conserved in Selaginella moellendorffii (lycophyte) and Physcomitrella patens (moss), although the former contains the GA signalling pathway, the latter does not. In the lycophyte, GA treatment promotes the outer wall development on microspores, whereas treatment with GA biosynthesis inhibitors disturbs its development. Contrary, in the moss, GAMYB homologue knockouts also produce abnormal spores that resemble Selaginella microspores treated with GA biosynthesis inhibitors and pollen grains of rice gamyb mutant. Moreover, the knockouts fail to develop male organs, instead ectopically forming female organs. Thus, before the establishment of the GA signalling pathway, basal land plants, including mosses, contained a GAMYB-based system for spore and sexual organ development. Subsequently, during the evolution from mosses to basal vascular plants including lycophytes, GA signalling might have merged to regulate this pre-existing GAMYB-based system. Plant male reproduction is controlled by gibberellin signalling via the transcription factor GAMYB. Here, deletion of GAMYB in a moss species lacking a gibberellin signalling pathway gives rise to abnormal spores, suggesting that primitive plants used GAMYB for the control of sexual organ development.