LEAF DEVELOPMENT IN ANGIOSPERMS

Mutations in CUC1 and CUC2 (for CUP-SHAPED COTYLEDON), which are newly identified genes of Arabidopsis, caused defects in the separation of cotyledons (embryonic organs), sepals, and stamens (floral organs) as well as in the formation of shoot apical meristems. These defects were most apparent in the double mutant. Phenotypes of the mutants suggest a common mechanism for separating adjacent organs within the same whorl in both embryos and flowers. We cloned the CUC2 gene and found that the encoded protein was homologous to the petunia NO APICAL MERISTEM (NAM) protein, which is thought to act in the development of embryos and flowers.

Petunia embryos carrying the no apical meristem (nam) mutation fail to develop a shoot apical meristem. Occasional shoots on nam- seedlings bear flowers that develop ten instead of five primordia in the second whorl. Double mutants with the homeotic gene green petals show that nam acts independently of organ identify in whorl 2 and now also affects primordium number in whorl 3. The nam gene was isolated by transposon tagging. The encoded protein shares a conserved N-terminal domain with several other proteins of unknown function and thus represents a novel class of proteins. Strikingly, nam mRNA accumulates in cells at the boundaries of meristems and primordia. These data indicate a role for nam in determining positions of meristems and primordia.

The shoot apical meristem is responsible for above-ground organ initiation in higher plants, accomplishing continuous organogenesis by maintaining a pool of undifferentiated cells and directing descendant cells toward organ formation. Normally, proliferation and differentiation are balanced, so that the structure and size of the shoot meristem is maintained. However, Arabidopsis plants homozygous for mutations at the CLAVATA1 (CLV1) locus accumulate excess undifferentiated cells. We describe the molecular cloning and expression pattern of the CLV1 gene. It encodes a putative receptor kinase, suggesting a role in signal transduction. The extracellular domain is composed of 21 tandem leucine-rich repeats that resemble leucine-rich repeats found in animal hormone receptors. We provide evidence that CLV1 expression in the inflorescence is specifically associated with meristematic activity.