The proper timing of flowering is critical for successful reproduction. The perception of the seasonal cues of daylength changes and exposure to cold influences flowering time in many plant species through the photoperiod and vernalization pathways, respectively. Here we show that a plant homeodomain (PHD) finger-containing protein, VIN3-LIKE 1 (VIL1), participates in both the photoperiod and vernalization pathways in Arabidopsis thaliana by regulating expression of the related floral repressors
FLOWERING LOCUS C (FLC) and FLOWERING LO-CUS M (FLM). In the vernalization pathway, VIL1, along with VERNALIZATION INSENSITIVE 3 (VIN3), is necessary for the modifications to FLC and FLM chromatin that are associated with an epigenetically silenced state and with acquisition of competence to flower. In addition, VIL1 regulates FLM independently of VIN3 in a photoperiod-dependent manner.Supplemental material is available at http://www.genesdev.org.Received September 13, 2006; revised version accepted October 13, 2006. The transition to flowering is a critical developmental event in the plant life cycle. In many plants, the perception of seasonal changes affects the timing of this transition. Two environmental cues that many plant species monitor for flowering time control are the prolonged cold of winter and changing day length; these cues promote flowering through the vernalization and photoperiod pathways, respectively. Most accessions of Arabidopsis thaliana exhibit a facultative response to these signals: In the absence of either environmental cue, flowering is delayed but nevertheless eventually occurs.In Arabidopsis, vernalization results in the mitotically stable repression of the potent floral repressor, FLOW-ERING LOCUS C (FLC) (Michaels and Amasino 1999;Sheldon et al. 1999). Vernalization-mediated repression of FLC is associated with histone modifications such as methylation of histone H3 Lys 9 (H3K9) and Lys 27 (H3K27) as well as H3 deacetylation (Bastow et al. 2004;Sung and Amasino 2004). A plant homeodomain (PHD) finger-containing protein, VERNALIZATION , is required for FLC repression by vernalization and the associated modifications to FLC chromatin (Sung and Amasino 2004). Here we show that a related PHD finger-containing protein, VIN3-LIKE 1 (VIL1), identified in a screen for proteins that interact with VIN3, cooperates with VIN3 in the vernalizationmediated repression of FLC. Furthermore, independently of VIN3 activity, VIL1 mediates the photoperiod-specific repression of another member of the FLC clade. Thus, VIL1 is involved in the regulation of flowering by two environmental-sensing pathways.
Results and DiscussionBecause VIN3 is essential for the vernalization response (Sung and Amasino 2004) and PHD finger-containing proteins are often members of multisubunit chromatin remodeling complexes (Bienz 2006), we searched for potential components of a VIN3 complex using the yeast two-hybrid system. This screen revealed two independent VIN3-interacting clones encoding C-terminal regions of At3g24440. In...