Organ formation in animals and plants relies on precise control of cell state transitions to turn stem cell daughters into fully differentiated cells. In plants, cells cannot rearrange due to shared cell walls. Thus, differentiation progression and the accompanying cell expansion must be tightly coordinated across tissues. PLETHORA (PLT) transcription factor gradients are unique in their ability to guide the progression of cell differentiation at different positions in the growing Arabidopsis thaliana root, which contrasts with well-described transcription factor gradients in animals specifying distinct cell fates within an essentially static context. To understand the output of the PLT gradient, we studied the gene set transcriptionally controlled by PLTs. Our work reveals how the PLT gradient can regulate cell state by region-specific induction of cell proliferation genes and repression of differentiation. Moreover, PLT targets include major patterning genes and autoregulatory feedback components, enforcing their role as master regulators of organ development.

Continuous formation of somatic tissues in plants requires functional stem cell niches where undifferentiated cells are maintained. In Arabidopsis thaliana, PLETHORA (PLT) and SCARECROW (SCR) genes are outputs of apicalbasal and radial patterning systems, and both are required for root stem cell specification and maintenance. The WUSCHEL-RELATED HOMEOBOX 5 (WOX5) gene is specifically expressed in and required for functions of a small group of root stem cell organizer cells, also called the quiescent center (QC). PLT and SCR are required for QC function, and their expression overlaps in the QC; however, how they specify the organizer has remained unknown. We show that PLT and SCR genetically and physically interact with plant-specific teosinte-branched cycloidea PCNA (TCP) transcription factors to specify the stem cell niche during embryogenesis and maintain organizer cells post-embryonically. PLT-TCP-SCR complexes converge on PLT-binding sites in the WOX5 promoter to induce expression.

Cyclin-dependent kinases (CDKs) play essential roles in coordinate control of cell cycle progression. Activation of CDKs requires interaction with specific cyclin partners and phosphorylation of their T-loops by CDK-activating kinases (CAKs). The Arabidopsis thaliana genome encodes four potential CAKs. CAK2At (CDKD;3) and CAK4At (CDKD;2) are closely related to the vertebrate CAK, CDK7/p40MO15; they interact with cyclin H and phosphorylate CDKs, as well as the C-terminal domain (CTD) of the largest subunit of RNA polymerase II. CAK1At (CDKF;1) shows cyclin H-independent CDK-kinase activity and can activate a heterologous CAK, Mcs6, in fission yeast. In Arabidopsis, CAK1At is a subunit of a protein complex of 130 kD, which phosphorylates the T-loop of CAK2At and CAK4At and activates the CTD-kinase activity of CAK4At in vitro and in root protoplasts. These results suggest that CAK1At is a novel CAK-activating kinase that modulates the activity of CAK2At and CAK4At, thereby controlling CDK activities and basal transcription in Arabidopsis

Activation of cyclin-dependent kinases (CDKs) requires phosphorylation of a threonine residue within the Tloop by a CDK-activating kinase (CAK). Here we isolated an Arabidopsis cDNA (CAK4At) whose predicted product shows a high similarity to vertebrate CDK7/p40 MO15 . Northern blot analysis showed that expressions of the four Arabidopsis CAKs (CAK1At^CAK4At) were not dependent on cell division. CAK2At-and CAK4At-immunoprecipitates of Arabidopsis crude extract phosphorylated CDK and the carboxy-terminal domain (CTD) of the largest subunit of RNA polymerase II with di¡erent preferences. These results suggest the existence of di¡erential mechanisms in Arabidopsis that control CDK and CTD phosphorylation by multiple CAKs. ß

Cyclin-dependent protein kinases (CDKs) play key roles in the progression of the cell cycle in eukaryotes. A CDK-activating kinase (CAK) catalyzes the phosphorylation of CDKs to activate their enzyme activity; thus, it is involved in activation of cell proliferation. In plants, two distinct classes of CAK have been identified; CDKD is functionally related to vertebrate-type CAKs, while CDKF is a plant-specific CAK having unique enzymatic characteristics. Recently, CDKF was shown to phosphorylate and activate CDKDs in Arabidopsis. This led to a proposal that CDKD and CDKF constitute a phosphorylation cascade that mediates environmental or hormonal signals to molecular machineries that control the cell cycle and transcription. In this review, we have summarized the biochemical features of plant CAKs and discussed the manner in which they diverge from animal and yeast orthologs. We have introduced several transgenic studies in which CAK genes were used as a tool to modify the CDK activity and to analyze cell division and differentiation during organ development.