Rice GH3 gene family: Regulators of growth and development

Auxin is an indispensable hormone throughout the lifetime of nearly all plant species. Several aspects of plant growth and development are rigidly governed by auxin, from micro to macro hierarchies; auxin also has a close relationship with plant-pathogen interactions. Undoubtedly, precise auxin levels are vitally important to plants, which have many effective mechanisms to maintain auxin homeostasis. One mechanism is conjugating amino acid to excessive indole-3-acetic acid (IAA; main form of auxin) through some GH3 family proteins to inactivate it. Our previous study demonstrated that GH3-2 mediated broad-spectrum resistance in rice (Oryza sativa L.) by suppressing pathogen-induced IAA accumulation and downregulating auxin signaling. Here, we further investigated the expression pattern of GH3-2 and other GH3 family paralogs in the life cycle of rice and presented the possible function of GH3-2 on rice root development by histochemical analysis of GH3-2 promoter:GUS reporter transgenic plants.Key words: auxin, GH3 gene, indole-3-acetic acid, Oryza sativa, rootThe phytohormone auxin regulates tropism and organ development and influences phyllotaxis, vascular canalization and root patterning by exerting its effect on cell division, elongation and differentiation in plants.^1,2 Indole-3-acetic acid (IAA) is the most widespread form of auxin in most plants. Supraoptimal or insufficient concentration of auxin will cause plants to exhibit abnormal phenotypes. 3-9 Auxin homeostasis is partly sustained by the GH3 gene family, a supervisor of the fluctuation of auxin. Most GH3 genes contain auxin-responsive cis-acting elements (AuxRE) in their promoter regions and react rapidly and transiently to auxin signaling.¹ Nineteen GH3 paralogs have been discovered in Arabidopsis.¹⁰ According to the phylogenetic relationship and acyl acid substrate preference, these genes are classified into three groups (I, II and III), which catalyze the formation of jasmonates, salicylic acid, 4-substituted benzoates or IAA acyl acid amido conjugates.^11,12 The rice GH3 gene family includes 13 paralogs, 4 belonging to group I (GH3-3, -5, -6 and -12) and 9 to group II (GH3-1, -2, -4, -7, -8, -9, -10, -11 and -13); group III GH3 is absent in rice.¹⁰ Rice GH3-1, -2, -8 and -13 paralogs have been biochemically confirmed to have IAA-amido synthetase activity by in vivo or in vitro assays.^6–9 It is believed that other GH3 group II paralogs in rice may also possess this enzymatic activity. But why does rice have such a functionally redundant group of GH3 proteins, which disobeys the economic principle? The explanation could be based on the different temporal and spatial expression of the genes encoding these proteins.