An Arabidopsis thaliana line that is mutant for the R2R3 MYB gene, AtMYB4, shows enhanced levels of sinapate esters in its leaves. The mutant line is more tolerant of UV‐B irradiation than wild type. The increase in sinapate ester accumulation in the mutant is associated with an enhanced expression of the gene encoding cinnamate 4‐hydroxylase, which appears to be the principal target of AtMYB4 and an effective rate limiting step in the synthesis of sinapate ester sunscreens. AtMYB4 expression is downregulated by exposure to UV‐B light, indicating that derepression is an important mechanism for acclimation to UV‐B in A.thaliana. The response of target genes to AtMYB4 repression is dose dependent, a feature that operates under physiological conditions to reinforce the silencing effect of AtMYB4 at high activity. AtMYB4 works as a repressor of target gene expression and includes a repression domain. It belongs to a novel group of plant R2R3 MYB proteins involved in transcriptional silencing. The balance between MYB activators and repressors on common target promoters may provide extra flexibility in transcriptional control.

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SummaryTranscription factors containing a conserved DNA-binding domain similar to that of the proto-oncogene c-myb have been identified in nearly all eukaryotes. MYB-related proteins from plants generally contain two related helix-turnhelix motifs, the R2 and R3 repeats. It was estimated that Arabidopsis thaliana contains more than 100 R2R3-MYB genes. The few cases where functional data are available suggest an important role of these genes in the regulation of secondary metabolism, the control of cell shape, disease resistance, and hormone responses. To determine the full regulatory potential of this large family of regulatory genes, a systematic search for the function of all genes of this family was initiated. Sequence data for more than 90 different A. thaliana R2R3-MYB genes have been obtained. Sequence comparison revealed conserved amino acid motifs shared by subgroups of R2R3-MYB genes in addition to the characteristic DNA-binding domain. No significant clustering of the genes was detected, although they are not uniformly distributed throughout the A. thaliana genome.

Stomatal pores located on the plant epidermis regulate CO(2) uptake for photosynthesis and the loss of water by transpiration. The opening and closing of the pore is mediated by turgor-driven volume changes of two surrounding guard cells. These highly specialized cells integrate internal signals and environmental stimuli to modulate stomatal aperture for plant survival under diverse conditions. Modulation of transcription and mRNA processing play important roles in controlling guard-cell activity, although the details of these levels of regulation remain mostly unknown. Here we report the characterization of AtMYB60, a R2R3-MYB gene of Arabidopsis, as the first transcription factor involved in the regulation of stomatal movements. AtMYB60 is specifically expressed in guard cells, and its expression is negatively modulated during drought. A null mutation in AtMYB60 results in the constitutive reduction of stomatal opening and in decreased wilting under water stress conditions. Transcript levels of a limited number of genes are altered in the mutant, and many of these genes are involved in the plant response to stress. Our data indicate that AtMYB60 is a transcriptional modulator of physiological responses in guard cells and open new possibilities to engineering stomatal activity to help plants survive desiccation.

The CCAAT box is one of the most common cis-elements present in eukaryotic promoters and is bound by the transcription factor NUCLEAR FACTOR Y (NF-Y). NF-Y is composed of three subunits, NF-YA, NF-YB, and NF-YC. Unlike animals and fungi, plants have significantly expanded the number of genes encoding NF-Y subunits. We provide a comprehensive classification of NF-Y genes, with a separation of closely related, but distinct, histone fold domain proteins. We additionally review recent experiments that have placed NF-Y at the center of many developmental stress-responsive processes in the plant lineage.