The theory that bioactive gibberellins (GAs) act as inhibitors of inhibitors of plant growth was based originally on the slender pea (Pisum sativum) mutant (genotype la cry-s), but the molecular nature of this mutant has remained obscure. Here we show that the genes LA and CRY encode DELLA proteins, previously characterized in other species (Arabidopsis [Arabidopsis thaliana] and several grasses) as repressors of growth, which are destabilized by GAs. Mutations la and cry-s encode nonfunctional proteins, accounting for the fact that la cry-s plants are extremely elongated, or slender. We use the la and cry-s mutations to show that in roots, DELLA proteins effectively promote the expression of GA synthesis genes, as well as inhibit elongation. We show also that one of the DELLAregulated genes is a second member of the pea GA 3-oxidase family, and that this gene appears to play a major role in pea roots.It is well known that primary root growth is strongly influenced by the plant hormone GA (Davies, 2004). For example, the application of bioactive GA to roots treated with the growth inhibitor ancymidol completely restored growth to that of the untreated plants (Tanimoto, 1991). Yaxley et al. (2001) established the importance of GAs for root growth in peas (Pisum sativum) by using a variety of GA-deficient mutant plants. In the na-1 mutant, for example, root GA 1 levels, and root elongation, were significantly reduced compared with wild-type plants, and when the GA 1 content was restored to wild-type levels, so too was root elongation.The GAs act by destabilizing the growth inhibitory DELLA proteins (Peng et al., 1997;Harberd et al., 1998;Silverstone et al., 2001;Alvey and Harberd, 2005). In other words, GA acts as an ''inhibitor of an inhibitor '' (Harberd et al., 1998). Interestingly, there is also evidence that DELLA proteins promote the biosynthesis of active GAs. For example, in the Arabidopsis (Arabidopsis thaliana) DELLA mutant rga, the expression of the biosynthesis gene GA4 is reduced, indicating that high DELLA protein levels are associated with an up-regulation of GA synthesis genes (Silverstone et al., 2001 (Silverstone et al., 1998;Gubler et al., 2002). However, the available evidence indicates greater redundancy in dicots compared with monocots (Ikeda et al., 2001;Thomas and Hedden, 2006). There have been five DELLA genes isolated from Arabidopsis (GAI, RGA, RGL1, RGL2, and RGL3), yet only one in rice (SLR1), barley (SLN1), and maize (Zea mays; D8; Peng et al., 1997Peng et al., , 1999Silverstone et al., 1998;Ikeda et al., 2001;Chandler et al., 2002;Gubler et al., 2002), with the possibility of another DELLA gene in maize (D9; accession no. ABI84225). It should be noted, however, that DELLAs have been studied in fewer dicot model species than in monocot species. To date, DELLA-encoding genes from pea have not been reported, even though observations on the slender phenotype of pea triggered the early suggestion that GA acts an inhibitor of an inhibitor (Brian, 1957). The elongated slender phenotype, con...