Memories of winter: vernalization and the competence to flower

ABSTRACT The promotion of flowering in response to a prolonged exposure to cold temperatures (i.e. winter) is a useful adaptation for plant species that flower in the spring. This promotion is known as vernalization and results in a permanent memory of cold exposure. While the physiology of vernalization has been extensively studied in many species, the molecular mechanism of vernalization remains largely unknown. Recent studies, however, have revealed some of the molecular events that create the requirement for vernalization. In Arabidopsis, naturally occurring late‐flowering ecotypes and plants containing late‐flowering mutations in the autonomous floral‐promotion pathway are relatively late flowering unless cold treated. The vernalization requirement of these late‐flowering ecotypes and autonomous‐pathway mutants is largely created by an upregulation of the floral inhibitor FLOWERING LOCUS C (FLC). After cold treatment, as imbibed seeds or young seedlings, FLC transcript levels are downregulated and remain low for the remainder of the plant's life, but return to high levels in the next generation. Plants containing a constitutively expressed 35S:FLC construct remain late flowering after cold treatment, indicating that FLC levels must be downregulated for vernalization to be effective. Thus the epigenetic downregulation of FLC appears to be a major target of the vernalization pathway and provides a molecular marker of the vernalized state.