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The nuclear SHL protein is composed of a N-terminal BAH domain and a C-terminal PHD finger. Both domains are found in transcriptional regulators and chromatin-modifying proteins. Arabidopsis plants over-expressing SHL showed earlier flowering and senescence phenotype. To identify SHL regulated genes, expression profiles of 35S::SHL plants were established with Affymetrix ATH1 microarrays. About 130 genes showed reduced transcript levels, and about 45 genes showed increased transcript levels in 35S:: SHL plants. The up-regulated genes included AGL20 and AGL9, which most likely cause the early flowering phenotype of 35S:: SHL plants. Late-flowering SHL-antisense lines showed reduced AGL20 mRNA levels, suggesting that AGL20 gene expression depends on the SHL protein. The stronger expression of senescence- and defence-related genes (such as DIN2, DIN11 and PR-1) is in line with the early senescence phenotype of SHL-over- expressing plants. SHL-down-regulated genes included stress response genes and the PSR3.2 gene (encoding a beta- glucosidase). SHL over-expression did not alter the tissue specificity of PSR3.2 gene expression, but resulted in reduced transcript levels in both shoots and roots. Plants with glucocorticoid-inducible SHL over-expression were established and used for expression profiling as well. A subset of genes was identified, which showed consistent changes in the inducible system and in plants with constitutive SHL over-expression
Detailed analysis of brassinosteroid (BR)-regulated genes can provide evidence of the molecular basis of BR effects. Classical techniques (such as subtractive cDNA cloning) as well as cDNA and oligonucleotide microarrays have been applied to identify genes which are upregulated or downregulated after BR treatment or are differently expressed in BR-deficient or -insensitive mutants compared with wild type plants. Genes encoding cell-wall-modifying enzymes, enzymes of the BR biosynthetic pathway, auxin response factors, and transcription factors are subject to BR regulation. Effects on several other metabolic pathways and interactions with other phytohormones have been reported as well, although some of these effects may depend on certain environmental conditions (for example, light/dark or stress), the developmental stage of the plants, and tissue types. The identification of components of the BR signal transduction pathway revealed different modes of transcriptional control in animals and plants. Steroid signaling in plants comprises the plasma membrane receptor kinases BRI1 and BAK1 and intracellular protein phosphorylations. Thus, BR signaling in plants is reminiscent of growth factor and TGF-beta signal transduction in animals. The phosphorylation cascade could be a basis of extensive signaling cross-talk and thereby explain the complexity of BR responses
Brassinosteroids (BRs) are steroidal plant hormones with important regulatory roles in various physiological processes, including growth, xylem differentiation, disease resistance, and stress tolerance. Several components of the BR signal transduction pathway have been identified. The extracellular domains of receptor kinases such as BRI1 perceive BRs and transduce the signal via intracellular kinase domains. Within the cell further kinases and phosphatases determine the phosphorylation status of transcription factors such as BES1 and BZR1. These factors mediate major BR effects. Studies of BR-regulated genes shed light on the molecular mode of BR action. Genes encoding cell-wall-modifying enzymes, enzymes of the BR biosynthetic pathway, transcription factors, and proteins involved in primary and secondary metabolism are subject to BR-regulation. Gene expression data also point at interactions with other phytohormones and a role of BR in stress responses. This article gives a survey of the BR-signaling pathway. Two BR-responsive genes, OPR3 and EXO, are described in detail
Background: The EXO (EXORDIUM) gene was identified as a potential mediator of brassinosteroid (BR)-promoted growth. It is part of a gene family with eight members in Arabidopsis. EXO gene expression is under control of BR, and EXO overexpression promotes shoot and root growth. In this study, the consequences of loss of EXO function are described. Results: The exo loss of function mutant showed diminished leaf and root growth and reduced biomass production. Light and scanning electron microscopy analyses revealed that impaired leaf growth is due to reduced cell expansion. Epidermis, palisade, and spongy parenchyma cells were smaller in comparison to the wild-type. The exo mutant showed reduced brassinolide-induced cotyledon and hypocotyl growth. In contrast, exo roots were significantly more sensitive to the inhibitory effect of synthetic brassinolide. Apart from reduced growth, exo did not show severe morphological abnormalities. Gene expression analyses of leaf material identified genes that showed robust EXO-dependent expression. Growth-related genes such as WAK1, EXP5, and KCS1, and genes involved in primary and secondary metabolism showed weaker expression in exo than in wild-type plants. However, the vast majority of BR-regulated genes were normally expressed in exo. HA- and GFP-tagged EXO proteins were targeted to the apoplast. Conclusion: The EXO gene is essential for cell expansion in leaves. Gene expression patterns and growth assays suggest that EXO mediates BR-induced leaf growth. However, EXO does not control BR-levels or BR-sensitivity in the shoot. EXO presumably is involved in a signalling process which coordinates BR-responses with environmental or developmental signals. The hypersensitivity of exo roots to BR suggests that EXO plays a diverse role in the control of BR responses in the root.