OsFY, a Homolog of AtFY, Encodes a Protein that Can Interact with OsFCA-γ in Rice （Oryza sativa L.）

FCA and FY are flowering time related genes involved in the autonomous flowering pathwayin Arabidopsis.FCA interacts with FY to regulate the alternative processing of FCA pre-mRNA.The FCA/FY interaction is also required for the regulation of FLC expression,a major floral repressor in Arabidopsis.However,it is not clear if the regulation of this autonomous flowering pathway is also present in monocotplants,such as rice.Recently,alternative RNA processing of OsFCA was observed in rice,which stronglysuggested the existence of an autonomous flowering pathway in rice.In this work,we cloned the cDNA ofthe autonomous flowering pathway gene OsFY from rice.The predicted OsFY protein contained a conserved7 WD-repeat region and at least two Pro-Pro-Leu-Pro motifs compared to Arabidopsis FY.The protein-protein interaction between OsFY and OsFCA-γ,the key feature of their gene function,was also demon-strated using the yeast two-hybrid system.The GenBank database search provided evidence of expressionfor other autonomous pathway gene homologs in rice.These results indicate that the autonomous floweringpathway is present in monocots,and the regulation through FY and FCA interaction is conserved betweenmonocots and dicots.     

Environmentally coordinated epigenetic silencing of FLC by protein and long noncoding RNA components

In Arabidopsis, the role of the vernalization pathway is to repress expression of a potent floral repressor, FLOWERING LOCUS C (FLC), after a sufficient period of winter cold has been perceived. Following winter, the lack of FLC expression allows unimpeded operation of the photoperiod pathway and hence rapid flowering of vernalized plants in spring via the activation of floral integrator genes. Molecular studies revealed that regulation of the key floral repressor, FLC, is under the control of the interplay between Trithorax group (TrxG)-mediated activation and Polycomb group (PcG)-mediated repression. On-off switch of genes by TrxG and PcG is an evolutionarily conserved mechanism to coordinate cellular identity in eukaryotes. Regulation of FLC by external cues provides an excellent model system to study mechanisms in which cell identity is influenced by environment. In this review, we discuss coordinated contributions by protein and long noncoding RNA components to this environmentally induced epigenetic switch of a developmental program in plants.     

Repression of the floral transition via histone H2B monoubiquitination

The Rad6-Bre1 complex monoubiquitinates histone H2B in target gene chromatin, and plays an important role in positively regulating gene expression in yeast. Here, we show that the Arabidopsis relatives of the yeast Rad6, ubiquitin-conjugating enzyme 1 (UBC1) and UBC2, redundantly mediate histone H2B monoubiquitination, and upregulate the expression of FLOWERING LOCUS C ( FLC ; a central flowering repressor in Arabidopsis) and FLC relatives, and also redundantly repress flowering, the developmental transition from a vegetative to a reproductive phase that is critical in the plant life cycle. Moreover, we have found that Arabidopsis relatives of the yeast Bre1, including HISTONE MONOUBIQUITINATION 1 (HUB1) and HUB2, also upregulate the expression of FLC and FLC relatives, and that HUB1 genetically interacts with UBC1 and UBC2 to repress the floral transition. These findings are consistent with a model in which HUB1 and HUB2 specifically interact with and direct UBC1 and UBC2 to monoubiquitinate H2B in developmental genes, and thus regulate developmental processes in plants.     

Chromatin regulation of flowering

The transition to flowering is a major developmental switch in the life cycle of plants. In Arabidopsis (Arabidopsis thaliana), chromatin mechanisms play critical roles in flowering-time regulation through the expression control of key flowering-regulatory genes. Various conserved chromatin modifiers, plant-specific factors, and long noncoding RNAs are involved in chromatin regulation of FLOWERING LOCUS C (FLC, a potent floral repressor). The well-studied FLC regulation has provided a paradigm for chromatin-based control of other developmental genes. In addition, chromatin modification plays an important role in the regulation of FLOWERING LOCUS T (FT, encoding florigen), which is widely conserved in angiosperm species. The chromatin mechanisms underlying FT regulation in Arabidopsis are likely involved in the regulation of FT relatives and, therefore, flowering-time control in other plants.     

Functional redundancy and new roles for genes of the autonomous floral-promotion pathway

The early-flowering habit of rapid-cycling accessions of Arabidopsis (Arabidopsis thaliana) is, in part, due to the genes of the autonomous floral-promotion pathway (AP). The AP promotes flowering by repressing expression of the floral inhibitor FLOWERING LOCUS C (FLC). AP mutants are therefore late flowering due to elevated levels of FLC, and this late-flowering phenotype is eliminated by loss-of-function mutations in FLC. To further investigate the role of the AP, we created a series of double mutants. In contrast to the phenotypes of single mutants, which are largely limited to delayed flowering, a subset of AP double mutants show a range of defects in growth and development. These phenotypes include reduced size, chlorophyll content, growth rate, and fertility. Unlike the effects of the AP on flowering time, these phenotypes are FLC independent. Recent work has also shown that two AP genes, FCA and FPA, are required for the repression and, in some cases, proper DNA methylation of two transposons. We show that similar effects are seen for all AP genes tested. Microarray analysis of gene expression in AP single and double mutants, however, suggests that the AP is not likely to play a broad role in the repression of gene expression through DNA methylation: very few of the genes that have been reported to be up-regulated in DNA methylation mutants are misexpressed in AP mutants. Together, these data indicate that the genes of the AP play important and sometimes functionally redundant roles in aspects of development in addition to flowering time.     

Genetic interactions of the Arabidopsis flowering time gene FCA, with genes regulating floral initiation

The genes controlling the timing of the transition from vegetative to reproductive growth are likely candidates for regulators of genes initiating floral development. We have investigated the interaction of one particular gene controlling flowering time, FCA, with the meristem identity-genes TERMINAL FLOWER 1 (TFL1), APETALA 1 (AP1) and LEAFY (LFY) and the floral repression gene EMBRYONIC FLOWER 1 (EMF1). Double mutant combinations were generated and the phenotypes characterized. The influence of strong and intermediate fca mutant alleles on the phenotype conferred by a 35S-LFY transgene was also analysed. The results support a model where FCA function promotes flowering in multiple pathways, one leading to activation of LFY and AP1, and another acting in parallel with LFY and AP1. Only the latter pathway is predicted to be non-functional in the intermediate fca-4 allele. The results are also consistent with AP1 and TFL1 negatively regulating FCA function. Combination of Columbia fca and emf1 mutant alleles confirmed that FCA is required for the early flowering of emf1. EMF1 and FCA are therefore likely to operate in different floral pathways.     

early in short days 4, a mutation in Arabidopsis that causes early flowering and reduces the mRNA abundance of the floral repressor FLC

The plant shoot is derived from the apical meristem, a group of stem cells formed during embryogenesis. Lateral organs form on the shoot of an adult plant from primordia that arise on the flanks of the shoot apical meristem. Environmental stimuli such as light, temperature and nutrient availability often influence the shape and identity of the organs that develop from these primordia. In particular, the transition from forming vegetative lateral organs to producing flowers often occurs in response to environmental cues. This transition requires increased expression in primordia of genes that confer floral identity, such as the Arabidopsis gene LEAFY. We describe a novel mutant, early in short days 4 (esd4), that dramatically accelerates the transition from vegetative growth to flowering in Arabidopsis: The effect of the mutation is strongest under short photoperiods, which delay flowering of Arabidopsis: The mutant has additional phenotypes, including premature termination of the shoot and an alteration of phyllotaxy along the stem, suggesting that ESD4 has a broader role in plant development. Genetic analysis indicates that ESD4 is most closely associated with the autonomous floral promotion pathway, one of the well-characterized pathways proposed to promote flowering of Arabidopsis: Furthermore, mRNA levels of a floral repressor (FLC), which acts within this pathway, are reduced by esd4, and the expression of flowering-time genes repressed by FLC is increased in the presence of the esd4 mutation. Although the reduction in FLC mRNA abundance is likely to contribute to the esd4 phenotype, our data suggest that esd4 also promotes flowering independently of FLC. The role of ESD4 in the regulation of flowering is discussed with reference to current models on the regulation of flowering in Arabidopsis.