SHORT-ROOT (SHR) and SCARECROW (SCR) are required for stem cell maintenance in the Arabidopsis (Arabidopsis thaliana) root meristem, ensuring its indeterminate growth. Mutation of SHR and SCR genes results in disorganization of the quiescent center and loss of stem cell activity, resulting in the cessation of root growth. This paper reports on the role of SHR and SCR in the development of leaves, which, in contrast to the root, have a determinate growth pattern and lack a persistent stem cell niche. Our results demonstrate that inhibition of leaf growth in shr and scr mutants is not a secondary effect of the compromised root development but is caused by an effect on cell division in the leaves: a reduced cell division rate and early exit of the proliferation phase. Consistent with the observed cell division phenotype, the expression of SHR and SCR genes in leaves is closely associated with cell division activity in most cell types. The increased cell cycle duration is due to a prolonged S-phase duration, which is mediated by up-regulation of cell cycle inhibitors known to restrain the activity of the transcription factor, E2Fa. Therefore, we conclude that, in contrast to their specific roles in cortex/endodermis differentiation and stem cell maintenance in the root, SHR and SCR primarily function as general regulators of cell proliferation in leaves.Stem cells are undifferentiated, totipotent cells that are able to duplicate themselves and to form offspring that differentiates into multiple cell types. They are situated in a microenvironment, the stem cell niche, where extracellular signals maintain stem cell division at low rates and prevent differentiation (Ohlstein et al., 2004;Li and Xie, 2005). In plants, the best studied stem cell niches are within the root and shoot apical meristems. There, stem cells produce somatic daughter cells that go on dividing and expanding, thereby forming the postembryonic tissues and organs that make up the body of the plant. It is the balance between stem cell maintenance within the meristem and differentiation of cells that exit the niche that facilitates indeterminate root and shoot growth.SHORT-ROOT (SHR) and SCARECROW (SCR) are members of the GRAS family of transcription factors (Pysh et al., 1999;Lee et al., 2008), required for stem cell maintenance in the root apical meristem. Mutation of SHR and SCR genes causes a disorganization of the quiescent center (QC) and loss of stem cell activity, resulting in the depletion of proliferating cells in the root meristem and, consequently, cessation of root growth. Essentially, loss of SHR/SCR function renders root growth determinate. Furthermore, shr and scr mutants lack longitudinal cell divisions that separate the cortex/endodermis initial daughter cells, resulting in only one ground tissue cell layer (Benfey et al