Plant brassinosteroid hormones

In animals, a large number of steroid hormones play important roles in numerous processes including reproduction and differentiation. The biologically active plant steroid brassinolide (BL) was first discovered in the pollen of western rape in 1979 (Grove et al., 1979). This finding suggested that BL is indispensable for plant growth and differentiation. To date, more than 50 BL analogs have been identified, and the group has been termed brassinosteroids (BRs) (Fujioka and Yokota, 2003). Brassinosteroids have several biological activities, such as inducing cell elongation when applied at very low concentrations. For this reason, soon after their discovery, they were suggested to be a sixth type of plant hormone; however, for years BRs were not considered true plant hormones. The turning point in BR research was the discovery of the Arabidopsis dwarf mutants det2 and cpd in 1996 (Li et al., 1996; Szekeres et al., 1996). These BR-deficient mutants were found to revert to the wild-type phenotype following BR treatment. Concurrent with the analysis of these mutants, an outline of the biosynthetic pathway of BRs was being elucidated through chemical analysis. Following the isolation of det2 and cpd, a great number of BR-deficient mutants were identified. The mutant genes were found to encode proteins that catalyze the conversion of plant steroids to BR precursors. Eventually, BRs were widely recognized as important plant hormones indispensable for growth and differentiation (Clouse and Sasse, 1998). In parallel, mutants that are insensitive to BRs were isolated (Clouse et al., 1996; Li et al., 1997) with phenotypes very similar to those of the BR-biosynthesis mutants. Investigations of these mutants revealed several mechanisms of BR perception and signal transduction (Bishop and Koncz, 2002; Clouse, 2002). This review describes findings on the effects of BRs on plant growth, BR biosynthesis and catabolism, and BR signal transduction.