Proper regulation of homeotic gene expression is critical for stem cell fate in both plants and animals. In Arabidopsis thaliana, the WUSCHEL (WUS)-RELATED HOMEOBOX 5 (WOX5) gene is specifically expressed in a group of root stem cell organizer cells called the quiescent center (QC) and plays a central role in QC specification. Here, we report that the SEUSS (SEU) protein, homologous to the animal LIM-domain binding (LDB) proteins, assembles a functional transcriptional complex that regulates WOX5 expression and QC specification. SEU is physically recruited to the WOX5 promoter by the master transcription factor SCARECROW. Subsequently, SEU physically recruits the SET domain methyltransferase SDG4 to the WOX5 promoter, thus activating WOX5 expression. Thus, analogous to its animal counterparts, SEU acts as a multi-adaptor protein that integrates the actions of genetic and epigenetic regulators into a concerted transcriptional program to control root stem cell organizer specification.
SHORTROOT (SHR) is essential for stem cell maintenance and radial patterning in Arabidopsis (Arabidopsis thaliana) roots, but how its expression is regulated is unknown. Here, we report that the Elongator complex, which consists of six subunits (ELP1 to ELP6), regulates the transcription of SHR. Depletion of Elongator drastically reduced SHR expression and led to defective root stem cell maintenance and radial patterning. The importance of the nuclear localization of Elongator for its functioning, together with the insensitivity of the elp1 mutant to the transcription elongation inhibitor 6-azauracil, and the direct interaction of the ELP4 subunit with the carboxyl-terminal domain of RNA polymerase II, support the notion that Elongator plays important roles in transcription elongation. Indeed, we found that ELP3 associates with the premessenger RNA of SHR and that mutation of Elongator reduces the enrichment of RNA polymerase II on the SHR gene body. Moreover, Elongator interacted in vivo with SUPPRESSOR OF Ty4, a well-established transcription elongation factor that is recruited to the SHR locus. Together, these results demonstrate that Elongator acts in concert with SUPPRESSOR OF Ty4 to regulate the transcription of SHR.
Cryptochrome 1 (CRY1) is an important light receptor essential for de-etiolation of Arabidopsis seedlings. However, its function in regulating plant architecture remains unclear. Here, we show that mutation in CRY1 resulted in increased branching of Arabidopsis plants . To investigate the underlying mechanism, we analyzed the expression profiles of branching-related genes and found that the mRNA levels of Phytochrome Interaction Factor 4 ( PIF4 ) and PIF5 are significantly increased in the cry1 mutant. Genetic analysis showed that the pif4 or pif4pif5 mutant is epistatic to the cry1 mutant, and overexpression of PIF4 conferred increased branching. Moreover, we demonstrated that PIF4 proteins physically associate with the G-box motif within the PIF4 promoter to form a self-activated transcriptional feedback loop, while CRY1 represses this process in response to blue light. Taken together, this study suggests that the CRY1–PIF4 module regulates gene expression via forming a regulatory loop and shoot branching in response to ambient light conditions.
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