The interface between the cell cycle and plant growth regulators: a mini review

The aim was to review knowledge about the interface betweenplant growth regulators and molecular checkpoints of the cell cycle. Atwhat level of biochemical regulation of the cell cycle do plant growthregulators interface? Are there different levels of interfacingdependent on the plant growth regulator involved? As a preamble totackling these questions, we overview the eukaryotic cell cycle withparticular emphasis on checkpoints that regulate the transition fromG0-G1-S-phase and G2-M. Cytokinins feature strongly as activators ofcell division in plants both in vivo and in vitro.Recent research has shown that zeatin treatment led to the up-regulationof CycD3 in Arabidopsis. This is a D-type cyclin showing stronghomology with vertebrate D cyclins which themselves are up-regulated byextracellular growth factors. Benzyladenine treatment can also shortenthe duration of S-phase through recruitment of latent origins of DNAreplication. Kinetin is involved in the phosphoregulation of the G2-Mcheckpoint; the major cyclin-dependent kinase (Cdk) at this checkpointhas recently been shown to be dephosphorylated as a result of cytokinintreatment, an effect which can also be mimicked by the fission yeastCdc25 phosphatase. Hence, a picture emerges of a cytokinin-inducedcontinuum of cell cycle activation through the up-regulation of a plantD-type cyclin at the G1 checkpoint and the phosphoregulation of the Cdkat the G2/M checkpoint. During S-phase, we argue for a link betweencytokinins and the proteins associated with replication origins.Gibberellic acid (GA) treatment induces internode elongation. Indeepwater rice, this response is mediated, at least partly, by aGA-induced up-regulation of a cyclin-Cdk at the G2-M checkpoint. Recentevidence has also linked abscisic acid to a cyclin-dependent kinaseinhibitor. These, so-called CKIs are negative regulators of Cdks whichfits with ABA's general role in growth inhibition; we await news ofethylene interactions. We highlight two instances of plant growthregulator-cell cycle interfacing during development, arguing for aninvolvement in microtubule orientation as a prerequisite to leafinitiation, and suggest a link between IAA and the activation of celldivisions in the pericycle required for lateral root initiation. A newD-type cyclin, recently discovered in Arabidopsis, may have akey role in this process. Finally, a model is presented which features ageneralised cyclin-Cdk checkpoint exhibiting various interfaces with theplant growth regulators.