In Arabidopsis ton 2 mutants, abnormalities of the cortical microtubular cytoskeleton, such as disorganization of the interphase microtubule array and lack of the preprophase band before mitosis, markedly affect cell shape and arrangement as well as overall plant morphology. We present the molecular isolation of the TON 2 gene, which is highly conserved in higher plants and has a vertebrate homolog of unknown function. It encodes a protein similar in its C-terminal part to B ؆ regulatory subunits of type 2A protein phosphatases (PP2As). We show that the TON2 protein interacts with an Arabidopsis type A subunit of PP2A in the yeast two-hybrid system and thus likely defines a novel subclass of PP2A subunits that are possibly involved in the control of cytoskeletal structures in plants.
INTRODUCTIONPlants possess unique features in many aspects of development compared with animals. At the cellular level, plants are characterized by specific features such as the presence of a pecto-cellulosic cell wall, continuous cytoplasmic connections through the plasmodesmata, and the lack of cell motility during morphogenesis (Kaplan and Hagemann, 1991). Reflecting these unique characteristics, the highly dynamic cortical arrays of microtubules (MTs) and actin filaments have adopted various specialized arrangements in plants. The organization of cortical arrays is coordinated tightly with other cellular events, and the cortical cytoskeleton, plasmalemma, and cell wall operate as a continuum (Wyatt and Carpita, 1993). The cortical cytoskeleton reorganizes steadily during all stages of the plant cell life and plays a crucial role in governing the orientation of both cell division and expansion (Cyr and Palevitz, 1995). In contrast to animals, fungi, and protists, in land plants the establishment of division planes depends on (1) the positioning of the preprophase band (PPB), a transient cortical ring of MTs that precisely foretells the position of the cell division plane at the G2/prophase transition, and (2) guidance of the phragmoplast to this predetermined cortical site during cytokinesis. During interphase, arrays of parallel MTs encircle the cell at the cortex. The positioning of MTs within these cortical arrays appears to involve both MT dynamicity (nucleation, growth/shrinkage, stabilization, and severing) and translocation, but the relative roles of these events and the proteins involved are unknown.To understand the molecular mechanisms underlying MT arrangements, mutants impaired in MT functions are essential, and a number of proteins involved in MT organization or dynamics have been identified through a genetic approach (Azimzadeh et al., 2001). These include the Arabidopsis botero1 mutant (which also is allelic to the fra2 mutant) (Zhong et al., 2001), which displays incorrect orientation of interphase cortical MTs and defects in anisotropic growth in root tip cells (Bichet et al., 2001). The FRA2 gene encodes a protein with high similarity to the sea urchin p60 subunit of katanin, which is known to be involved in MT...