Particular significance of SRD2-dependent snRNA accumulation in polarized pattern generation during lateral root development of Arabidopsis.

Lateral root primordia are initiated by anticlinal division of cells in the pericycle and are constructed through an ordered set of cell divisions. At the completion of the development of the primordium, cell division ceases, after which the lateral root meristem is activated. In Arabidopsis, this course of lateral root morphogenesis was found to be significantly susceptible to srd2-1, a temperature-sensitive mutation of the SRD2 gene encoding an activator of small nuclear RNA (snRNA) transcription. The srd2-1 mutation altered the organization of cells of the root primordium and, importantly, maintained primordial cell division for a long period, resulting in the formation of abnormal hemispherical laterals. Expression patterns of various reporter genes suggested that both the apical-basal and radial axes were not well established in the lateral root primordia of the srd2-1 mutant. In the early stages of development of the primordium, the srd2-1 mutation reduced the amount of the auxin efflux facilitator PIN-FORMED (PIN) and, probably by this means, interfered with the generation of an auxin gradient. Spliceosomal snRNAs accumulated at high levels throughout young root primordia and then decreased in association with the arrest of cell division, and finally increased again when the apical meristem became activated. The accumulation of snRNAs was severely suppressed by the srd2-1 mutation, and this was detectable before any morphological defect became evident. These findings suggest that high-level accumulation of snRNA involving the SRD2 function is particularly important for expression of PINs in polarized pattern generation during the development of lateral root primordia.