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Common mechanisms regulating expression of rice aleurone genes ...

Common mechanisms regulating expression of rice aleurone genes ...

Common mechanisms regulating expression of rice aleurone genes

Biochimica et Biophysica Acta 1759 (2006) 478–490 www.elsevier.com/locate/bbaexp Promoter paper Common mechanisms regulating expression of rice aleurone genes that contribute to the primary response for gibberellin Kenji Washio ⁎ , Masaaki Morikawa Group of Environmental Molecular Biology, Section of Environmental Bioscience, Faculty of Environmental Earth Science, Hokkaido University, Sapporo 060-0810, Japan Received 5 April 2006; received in revised form 7 September 2006; accepted 7 September 2006 Available online 16 September 2006 Abstract During germination of cereal seeds, aleurone cells respond to gibberellins (GA) by synthesizing and secreting hydrolytic enzymes that mobilize the reserved nutrients. It has been shown that products of early GA response genes, like a transcription factor GAMyb, act as key molecules leading to this regulation. Pivotal roles of GAMyb on expression of hydrolase genes have been well documented, whereas regulation of GAMyb expression itself remains obscure. In order to understand virtual mechanisms of the GA-mediated expression of genes, it is important to know how GA control expression of early GA response genes. Using an aleurone transient expression system of rice (Oryza sativa L.), we examined GA responsive domains of early GA response genes in the aleurone, such as GAMyb and OsDof3. The upstream promoter could not confer GA response. Extensive analyses revealed the presence of enhancer-like activities in a large first intron. In Arabidopsis, intron enhancers have been identified in MADS-box homeotic genes, AGAMOUS (AG) and FLOWERING LOCUS C (FLC), in which large introns should not only confer proper gene expressions, but also associate with gene silencing by covalent modifications of both DNA and histone. These evidences prompt us to assign that chromatin-based control might be important for initial GA action. Based on this assumption, we have identified DNA methylation of the GAMyb locus in germinated rice seeds. © 2006 Elsevier B.V. All rights reserved. Keywords: Aleurone; DNA methylation; GAMyb; Gibberellin; Rice; Seed germination 1. Introduction Gibberellins (GA) are tetracyclic diterpenoid hormones essential for many aspects of plant growth and development, such as seed germination, stem elongation, leaf expansion, flower initiation and anther development [1,2]. Molecular genetic analyses of the GA-related mutants have identified crucial components of GA signaling [3,4]. The DELLA proteins, a subgroup of the GRAS family of putative transcription factors [5], are located in the nucleus and rapidly disappeared after GA application [6–8]. Loss-of-function mutations in the DELLA gene produce a constitutive GA response phenotype in some cases [9,10]. Expression of the DELLA protein lacking the DELLA domain, which is a constitutive repressor form, shows a dominant GA-insensitive ⁎ Corresponding author. Tel.: +81 11 706 2378; fax: +81 11 706 2253. E-mail address: washi@ees.hokudai.ac.jp (K. Washio). phenotype [9–12]. These results provide a reliable concept that the DELLA proteins are robust repressors of GA signaling and the functions of GA are established by removing the repressions of the DELLA proteins [6,13]. Other GA signaling components illustrate molecular events that involve the degradation of the DELLA proteins. The F-box protein is subunit of a SCF E3 ubiquitin ligase complex that plays a key role for substrate specificities in the 26S proteasome pathway [14]. Mutations in both Arabidopsis SLY1 [15] and rice GID2 [16], encoding GA-related F-box proteins, result in a GA-insensitive dwarfism. In these mutants, DELLA proteins accumulate at high-levels in nuclei even in the presence of GA. These findings indicate that this F-box protein class is positive regulators of GA signaling and capable of destabilizing the DELLA proteins via proteolysis. Recently, an initial factor of GA signaling has been proposed. Ueguchi-Tanaka et al. identified a recessive rice mutant gid1 that showed a strong GA-insensitive dwarfism [17]. Mutant phenotypes suggest that 0167-4781/$ - see front matter © 2006 Elsevier B.V. All rights reserved. doi:10.1016/j.bbaexp.2006.09.001

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