During early seed development, nuclear divisions in the endosperm are not followed by cell division, leading to the development of a syncytium. The simple organization of the Arabidopsis endosperm provides a model in which to study the regulation of the cell cycle in relation to development. To monitor nuclear divisions, we constructed a HISTONE 2B::YELLOW FLUORESCENT PROTEIN gene fusion ( H2B::YFP ) . To validate its use as a vital marker for chromatin in plants, H2B::YFP was expressed constitutively in Arabidopsis. This enabled the observation of mitoses in living root meristems. H2B::YFP was expressed specifically in Arabidopsis syncytial endosperm by using GAL4 transactivation. Monitoring mitotic activity in living syncytial endosperm showed that the syncytium was organized into three domains in which nuclei divide simultaneously with a specific time course. Each mitotic domain has a distinct spatiotemporal pattern of mitotic CYCLIN B1;1 accumulation. The polar spatial organization of the three mitotic domains suggests interactions between developmental mechanisms and the regulation of the cell cycle.
INTRODUCTIONRegulation of the cell cycle in plants and animals involves a remarkable number of conserved genes and mechanisms (Huntley and Murray, 1999;Mironov et al., 1999). In animals, regulation of the entry into G1 has proven to be critical for some developmental steps, such as Drosophila wing patterning (Johnston and Edgar, 1998) and the integration between trophic factors and proliferation (Conlon and Raff, 1999;Galloni and Edgar, 1999). Recently, an essential block of entry into mitosis has been characterized for the coordination between proliferation and gastrulation in Drosophila (Grosshans and Wieschaus, 2000;Mata et al., 2000;Seher and Leptin, 2000). It has been shown in plants that endogenous cell cycle regulation is very precise in meristems, where cells are produced in roots (Berger et al., 1998a), and in the shoot apex (Meyerowitz, 1997;Laufs et al., 1998). However, little is known about the general mechanisms that govern cell proliferation in relation to patterning in plants. To undertake such an analysis, we propose to use a simple developmental system, the syncytial endosperm in Arabidopsis.Several developmental programs in multicellular organisms are characterized by multiple nucleate structures called syncytia or coenocytes. These structures originate either from the fusion of multiple cells, as in osteoclasts (Jotereau and Le Douarin, 1978), or from a single cell in which nuclear divisions proceed without cytokinesis. The best-studied examples of the latter type are the Drosophila oocyte (Foe et al., 1993) and the hyphae of the fungus Aspergillus nidulans (Doonan, 1992). In these systems, nuclei divide synchronously and thus provide simple models in which to study the developmental control of the cell cycle (Edgar and Lehner, 1996). In plants, the development of syncytia is typical of the endosperm (Vijayaraghavan and Prabhakar, 1984;Lopes and Larkins, 1993; Berger, 1999). Endosperm is inc...