We describe two recessive alleles of a rice heterochronic gene, plastochron1-1 ( pla1-1 ) and pla1-2 , that reduce the length of the plastochron to approximately half that of the wild type. Because the onset of the reproductive phase in pla1 was not temporally affected, the number of leaves produced in the vegetative phase was nearly twice that produced in the wild type. Panicle development was severely disturbed in pla1 mutants. In pla1-1 , many primordia of primary rachis branches were converted into vegetative shoots. These ectopic shoots repeated the initiation of panicle development and the conversion of primary rachis branches into shoots. In the weak allele pla1-2 , however, only the basal one or two primordia developed as vegetative shoots, and the remaining primordia developed to produce a truncated panicle. These results indicate that both vegetative and reproductive programs are expressed simultaneously during the reproductive phase of pla1 ; however, the degree varied depending on the strength of the allele. Accordingly, pla1 is a heterochronic mutation that extends the vegetative period. The shoot apical meristem of pla1 was larger than that of the wild type, although the shape was not modified. An in situ hybridization experiment using the histone H4 gene as a probe revealed that cell divisions are accelerated in the pla1 meristem. The PLA1 gene is considered to regulate the duration of the vegetative phase by controlling the rate of leaf production in the meristem.
INTRODUCTIONPlant development generally is divided into four distinct phases: embryogenesis, early vegetative phase (juvenile phase), late vegetative phase (adult phase), and reproductive phase (Conway and Poethig, 1993). In almost all plant species, two phase changes are recognized easily by the distinct morphological changes that occur in plants from the embryonic phase to the juvenile vegetative phase and from the adult vegetative phase to the reproductive phase. A clear change between the juvenile and adult vegetative phases has been reported for English ivy, maize, and Arabidopsis (Hackett, 1985;Zimmerman et al., 1985;Poethig, 1988Poethig, , 1990 Lawson and Poethig, 1995;Telfer et al., 1997). However, in several species including rice, the phase change from juvenile to adult is not clear because it is continuous and is not accompanied by gross visible changes.Whatever the case may be, the plant life cycle is driven by phase-specific genetic programs that are partially overlapping (Bongard-Pierce et al., 1996). Mutations that delay or advance phase change have been reported for several plant species. Late-flowering mutants can be interpreted as prolonging the vegetative phase and delaying the reproductive phase. Recently, using trichome distribution or epidermal characters as phase-specific markers, researchers have described several late-flowering and dwarf mutations that prolong the juvenile phase (Evans and Poethig, 1995;Telfer et al., 1997).Heterochronic mutations affecting the timing of developmental events may be of major sig...