The interplay among histone modifications modulates the expression of master regulatory genes in development. Chromatin effector proteins bind histone modifications and translate the epigenetic status into gene expression patterns that control development. Here, we show that two Arabidopsis thaliana paralogs encoding plant-specific proteins with a plant homeodomain (PHD) motif, SHORT LIFE (SHL) and EARLY BOLTING IN SHORT DAYS (EBS), function in the chromatin-mediated repression of floral initiation and play independent roles in the control of genes regulating flowering. Previous results showed that repression of the floral integrator FLOWERING LOCUS T (FT) requires EBS. We establish that SHL is necessary to negatively regulate the expression of SUPPRESSOR OF OVEREXPRESSION OF CO1 (SOC1), another floral integrator. SHL and EBS recognize di-and trimethylated histone H3 at lysine 4 and bind regulatory regions of SOC1 and FT, respectively. These PHD proteins maintain an inactive chromatin conformation in SOC1 and FT by preventing high levels of H3 acetylation, bind HISTONE DEACETYLASE6, and play a central role in regulating flowering time. SHL and EBS are widely conserved in plants but are absent in other eukaryotes, suggesting that the regulatory module mediated by these proteins could represent a distinct mechanism for gene expression control in plants.
The rotation of the earth results in periodic changes in environmental factors such as daylength and temperature; the circadian clock is the endogenous mechanism responsible for day-length measurement, and allows plants to anticipate these fluctuations and modulate their developmental programs to maximize adaptation to those environmental cues. Flowering represents the transition from a vegetative to reproductive phase and is controlled by complex and highly regulated genetic pathways. In many plants, the time of flowering is strongly influenced by photoperiod, which synchronizes the floral transition with the favourable season of the year. Over the last decade, genetic approaches have aided the discovery of many signalling components involved in the photoperiod pathway and here, we highlight the significant progress made in identifying the molecular mechanisms that measure daylength and control flowering initiation in Arabidopsis, a long day (LD) plant, and in rice, a short day (SD) plant. Some components of the Arabidopsis regulatory network are conserved in other species, but the difference in the function of particular genes may contribute to the opposite photoperiodic flowering response observed between LD and SD plants. The specific regulatory mechanisms involved in controlling CONSTANS (CO) expression and stability by the circadian clock and the different photoreceptors will be described. In addition, the role of FLOWERING LOCUS T (FT), as part of the florigen, and several other light signalling and circadian-dependent components in photoperiodic flowering will be also discussed.Additional key words: circadian clock, CONSTANS, florigen, FT, photoperiodism, photoreceptors. Resumen Revisión. Control fotoperiódico del tiempo de floraciónLa rotación diaria de la tierra provoca cambios periódicos en la duración del día o en la temperatura, y el reloj circadiano es un mecanismo endógeno responsable de la medida de la duración del día; este oscilador molecular permite a los organismos anticiparse a dichos cambios y adaptar su desarrollo de manera adecuada. La floración representa la transición desde una fase vegetativa del crecimiento a una reproductiva, y está controlada por diferentes rutas, muy complejas y altamente reguladas. En muchas plantas, esta transición está controlada principalmente por la duración del día o fotoperiodo, el cual sincroniza la floración con la estación más favorable del año. En la última década, diferentes estudios genéticos han facilitado la caracterización de muchos componentes de señalización involucrados en la ruta del fotoperiodo y en esta revisión se discute el progreso reciente que se ha llevado a cabo en la identificación de los mecanismos moleculares que permiten medir la duración del día, y que controlan el tiempo de floración en Arabidopsis, una especie de día largo, y en arroz, un especie de día corto. Aunque los componentes de las rutas reguladoras de Arabidopsis parecen conservarse en otras especies, la diferencia de función de genes concretos parece contribuir a...
The Arabidopsis protein EARLY BOLTING IN SHORT DAYS (EBS), a plant-specific transcriptional regulator, is involved in the control of flowering time by repressing the floral integrator FT. The EBS protein binds the H3K4me3 histone mark and interacts with histone deacetylases to modulate gene expression. Here, we show that EBS also participates in the regulation of seed dormancy. ebs mutations cause a reduction in seed dormancy, and the concurrent loss of function of the EBS homologue SHORT LIFE (SHL) enhances this dormancy alteration. Transcriptomic analyses in ebs mutant seeds uncovered the misregulation of several regulators of seed dormancy including the MADS box gene AGAMOUS-LIKE67 (AGL67). AGL67 interacts genetically with EBS in seed dormancy regulation, indicating that both loci act in the same pathway. Interestingly, EBS functions independently of the master regulator gene of dormancy DELAY OF GERMINATION 1 (DOG1) and other genes encoding chromatin remodelling factors involved in the control of seed dormancy. Altogether, these data show that EBS is a central repressor of germination during seed dormancy and that SHL acts redundantly with EBS in the control of this developmental process. Our observations suggest that a tightly regulated crosstalk among histone modifications is necessary for a proper control of seed dormancy.
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