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3. | Dwarfism and reduced responses to gibberellic acid in transgenic rice plants expressing the Arabidopsis GAI gene, and the dominant negative allele gai |
X. FU, D.E. RICHARDs, N.P. HARBERD and P. CHRISTOU John Innes Center, Colney Lane, Norwich NR4 7UH, UK |
Gibberellins (GAs) are tetracyclic diterpenoid hormones essential for
plant growth and development. Mutants deficient for GA synthesis, perception
and signal transduction have been isolated from a number of species, and
the analysis of GA mutants in Arabidopsis has significantly advanced
knowledge of the GA signaling pathway (Richards et al. 2000). Isolation
of the Arabidopsis gai (gibberellic acid insensitive) mutant led
to the cloning of the GAI gene (Peng et al. 1997). This
encodes a transcription factor of the GRAS family, named after its three
principal members: GAI, RGA and SCARECROW. The gai mutation is
dominant to the wild type allele, and similar mutations have been identified
in wheat and maize, causing genetically dominant dwarfism and reductions
in the response to GA (Richards et al. 2000). Surprisingly, high level expression of the wild type allele, GAI, also causes dwarfism
Since the dominant loss of function allele gai causes dwarfism and a reduced GA growth response in rice, it may seem surprising that overexpressing the wild type allele GAI has a similar effect. Furthermore, the phenotype of the gai transgene cannot be rescued by exogenous GA, whereas that of the GAI transgene can be rescued as long as it is expressed at low levels. These results confirm a derepressible repressor model, in which the normal function of the GAI protein is to suppress plant growth. In normal development, the activity of GAI is antagonized by GA. In plants where GAI protein is expressed as a transgene, the increased activity can be suppressed by increasing the doses of GA, at least up to a certain point. High level expression of GAI is resistant to GA, perhaps because the GA signaling pathway becomes saturated and there remains an excess of the active GAI protein. Presumably, the dominant gai allele encodes a repressor that cannot be antagonized by GA, so that even relatively low levels of the protein succeed in stunting plant growth (Richards et al. 2000). These results indicate that the mechanism of GAI activity has been conserved over millions of years of plant evolution. The dominant GA-resistant allele from Arabidopsis (gai) can impose its repressive activity on the growth response genes in rice, and high level expression of the wild type allele has a similar effect. The resulting dwarf phenotypes are similar to those seen in the maize and wheat 'green revolution' genotypes (Peng et al. 1999), and it is hoped that similar increases in grain yield, at the expense of straw biomass, can be achieved in rice. References Peng, J.R., P. Carol, D.E. Richards, K.E. King, R.J. Cowling, G.P. Murphy and N.P. Harberd, 1997. The Arabidopsis GAI gene defines a signaling pathway that negatively regulates gibberellin responses. Genes & Dev 11: 3194-3205. Peng, J.R., D.E. Richards, N.M. Hartley, G.P. Murphy, K.M. Devos, J.E. Flitham, J. Beales, L.J. Fish, A.J. Worland, F. Pelica, D. Sudhakar, P. Christou, J.W. Snape, M.D. Gale and N.P. Harberd, 1999. "Green revolution" genes encode mutant gibberellin response modulators. Nature 400: 256-261. Richards, D.E., J.R. Peng and N.P. Harberd, 2000. Plant GRAS and metazoan STATs: one family? Bioessays 22: 573-577. |
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