Organogenesis at the shoot meristem requires a delicate balance between stem cell specification and differentiation. In Arabidopsis thaliana, WUSCHEL (WUS) is a key factor promoting stem cell identity, whereas the CLAVATA (CLV1, CLV2, and CLV3) loci appear to promote differentiation by repressing WUS expression. In a screen for mutations modifying clv1 mutants, we have identified a novel regulator of meristem development we term CORONA (CNA). Whereas cna single mutant plants exhibit subtle defects in meristem development, clv cna double mutants develop massively enlarged apices that display early loss of organogenesis, misexpression of WUS and CLV3, and eventual differentiation of the entire apex. The CNA gene was isolated by positional cloning and found to encode a class III homeodomain Leu zipper protein. A missense mutation resulting in the dominant-negative cna-1 allele was identified in a conserved domain of unknown function, and a likely null allele was shown to display a similar but weaker phenotype. CNA is expressed in developing vascular tissue, diffusely through shoot and flower meristems, and within developing stamens and carpels. Our analysis of WUS expression in wild-type, clv, and clv cna plants revealed that, contrary to current models, WUS is neither necessary nor sufficient for stem cell specification and that neither WUS nor CLV3 is a marker for stem cell identity. We propose that CNA functions in parallel to the CLV loci to promote organ formation.

No abstract

A central feature of plant development is the continuous generation of organs throughout the lifespan of the plant. In the region designated the meristem, undifferentiated cells are maintained throughout the life cycle of the plant, providing a source of cells from which plant organs are derived. Cells that are differentiating as they are channelled toward a particular developmental fate also reside in the meristem, as do the resultant organ primordia. The meristem is a self‐renewing structure and its stem cell population is maintained at a near constant number despite the perpetual mobilisation of differentiating cells into organogenesis. The meristem's capacity to balance continuous differentiation of cells while replenishing the pool of undifferentiated, pluripotent cells is tightly controlled by a very complex and overlapping network of regulatory pathways. Key Concepts: Plants, unlike animals, have the capacity to generate new organs post‐embryonically, throughout the lifespan of the plant. Shoot and root apical meristems have the capacity to balance perpetual differentiation of cells while maintaining a population of undifferentiated stem cells. Because fates of meristematic cells are determined by their relative positions, they must be in continuous communication with their neighbouring cells. Signals can be transmitted from more mature cells to initial cells to specify the pattern of meristem differentiation. The number of stem cells in meristems is remarkably constant despite the continuous recruitment of differentiating cells into organs. In Arabidopsis shoot meristems, a feedback loop between WUSCHEL and the CLAVATA signalling pathway is crucial for specifying and maintaining the stem cell niche in the shoot apex. Within their niche boundaries, stem cells remain in an undifferentiated state in response to positional cues from neighbouring cells while cells displaced from the niche begin the process of differentiation.

In the region designated the shoot meristem, undifferentiated cells are maintained throughout the life cycle of the plant, providing a source of cells from which plant organs are derived. Cells that are differentiating as they are channelled toward a particular developmental fate also reside in the shoot meristem, as do the resultant organ primordia.