e-journal
Transcription factor HAT1 is phosphorylated by BIN2 kinase and mediates brassinosteroid repressed gene expression in Arabidopsis
Plant steroid hormones, brassinosteroids (BRs), play essential roles in modulating cell elongation, vascular
differentiation, senescence and stress responses. BRs signal through plasma membrane-localized receptor
and other components to modulate the BES1/BZR1 (BRI1-EMS SUPPRESSOR 1/BRASSINAZOLE RESISTANT
1) family of transcription factors that modulate thousands of target genes. Arabodopsis thaliana homeodomain-
leucine zipper protein 1 (HAT1), which encodes a homeodomain-leucine zipper (HD-Zip) class II transcription
factor, was identified through chromatin immunoprecipitation (ChIP) experiments as a direct
target gene of BES1. Loss-of-function and gain-of-function mutants of HAT1 display altered BR responses.
HAT1 and its close homolog HAT3 act redundantly, as the double mutant hat1 hat3 displayed a reduced BR
response that is stronger than the single mutants alone. Moreover, hat1 hat3 enhanced the phenotype of a
weak allele of the BR receptor mutant bri1 and suppressed the phenotype of constitutive BR response
mutant bes1-D. These results suggest that HAT1 and HAT3 function to activate BR-mediated growth.
Expression levels of several BR-repressed genes are increased in hat1 hat3 and reduced in HAT1OX, suggesting
that HAT1 functions to repress the expression of a subset of BR target genes. HAT1 and BES1 bind
to a conserved homeodomain binding (HB) site and BR response element (BRRE) respectively, in the promoters
of some BR-repressed genes. BES1 and HAT1 interact with each other and cooperate to inhibit BRrepressed
gene expression. Furthermore, HAT1 can be phosphorylated and stabilized by GSK3 (GLYCOGEN
SYNTHASE KINASE 3)-like kinase BIN2 (BRASSINOSTEROID-INSENSITIVE 2), a well established negative
regulator of the BR pathway. Our results thus revealed a previously unknown mechanism by which BR
signaling modulates BR-repressed gene expression and coordinates plant growth.
Keywords: brassinosteroids, BES1, HAT1, HAT3, transcription repression, BIN2, phosphorylation, Arabidopsis
thaliana.
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