The chronic food shortage that was feared after the rapid expansion of the world population in the 1960s was averted largely by the development of a high-yielding semi-dwarf variety of rice known as IR8, the so-called rice 'green revolution'. The short stature of IR8 is due to a mutation in the plant's sd1 gene, and here we identify this gene as encoding an oxidase enzyme involved in the biosynthesis of gibberellin, a plant growth hormone. Gibberellin is also implicated in green-revolution varieties of wheat, but the reduced height of those crops is conferred by defects in the hormone's signalling pathway.

Gibberellin (GA) regulates growth and development in plants. We isolated and characterized a rice GA-insensitive dwarf mutant, gid2. The GID2 gene encodes a putative F-box protein, which interacted with the rice Skp1 homolog in a yeast two-hybrid assay. In gid2, a repressor for GA signaling, SLR1, was highly accumulated in a phosphorylated form and GA increased its concentration, whereas SLR1 was rapidly degraded by GA through ubiquitination in the wild type. We conclude that GID2 is a positive regulator of GA signaling and that regulated degradation of SLR1 is initiated through GA-dependent phosphorylation and finalized by an SCF(GID2)-proteasome pathway.

SummaryThe phytohormone gibberellin (GA) controls growth and development in plants. Previously, we identi®ed a rice F-box protein, gibberellin-insensitive dwarf2 (GID2), which is essential for GA-mediated DELLA protein degradation. In this study, we analyzed the biological and molecular biological properties of GID2. Expression of GID2 preferentially occurred in rice organs actively synthesizing GA. Domain analysis of GID2 revealed that the C-terminal regions were essential for the GID2 function, but not the N-terminal region. Yeast two-hybrid assay and immunoprecipitation experiments demonstrated that GID2 is a component of the SCF complex through an interaction with a rice ASK1 homolog, OsSkp15. Furthermore, an in vitro pulldown assay revealed that GID2 speci®cally interacted with the phosphorylated Slender Rice 1 (SLR1). Taken these results together, we conclude that the phosphorylated SLR1 is caught by the SCF GID2 complex through an interacting af®nity between GID2 and phosphorylated SLR1, triggering the ubiquitin-mediated degradation of SLR1.

A rice semi-dwarf variety, IR8, known as "miracle rice" enabled dramatic increases rice production and its widespread adoption averted predicted food shortages in Asia during the 1960s to 1990s. This remarkable achievement was referred to as "green revolution". The short stature of IR8 was derived from the semi-dwarf gene, sd1, and the sd1 gene contributed significantly to the rice "green revolution". In this paper, we described the physiological, molecular genetic and biochemical characterization of the sd1 gene. The sd1 mutant contained lower gibberellin (GA) levels than wild-type plants but responded sensitively to exogenous GA. Cloning and sequence analyses revealed that the SD1 gene encoded a GA biosynthetic enzyme, GA20 oxidase. In all of the sd1 mutants tested, nucleotide deletions or substitutions were observed in the GA20 oxidase gene (GA20ox-2), which induced an internal stop codon or single amino acid substitutions, respectively. The sd1 plants, which the wild-type GA20ox-2 gene was introduced showed the normal height. A recombinant GA20ox-2 protein produced from the cDNA clone in E. coli catalyzed the conversion of GA 53 to GA 20 . These results confirmed that SD1 encodes an active GA20 oxidase. The expression of GA20ox-2 was down-regulated by GA in a similar manner to that of some GA20oxs in other plants. The rice genome carried at least two GA 20-oxidase genes (GA20ox-1 and GA20ox-2) and SD1 corresponded to GA20ox-2, which is highly expressed in the leaves and flowers, whereas GA20ox-1 is preferentially expressed in the flowers. The reduced plant height associated with the sd1 alleles was due to the low amount of active GA in leaves, which was caused by a mutation of the GA20ox-2 gene. On the basis of these results, we discussed the importance of GA in the regulation of plant height in crop breeding.