Epigenetic inheritance and reprogramming in plants and fission yeast.

Plants and fission yeast exhibit a wealth of epigenetic phenomena, including transposon regulation, heterochromatic silencing, and gene imprinting. They provide excellent model organisms to address the question of how epigenetic information is propagated to daughter cells. We have addressed the questions of establishment, maintenance, and inheritance of heterochromatic silencing using the fission yeast Schizosaccharomyces pombe and the plant Arabidopsis thaliana by using a variety of genetic and genomic approaches. We present here results showing the cell cycle dependence of RNA in fission yeast RNA interference (RNAi), which is required for proper transcriptional silencing of the centromeric heterochromatin, and that this process occurs during S phase, allowing for precise copying and reestablishment of heterochromatic histone modifications following DNA replication and cell division. We also show that in plants, cells in culture and male germ-line cells undergo massive epigenomic changes correlated with the appearance of a novel class of 21-nucleotide small interfering RNA (siRNA) from transcriptionally reactivated transposable elements (TEs) following loss of heterochromatic DNA and histone methylation. We propose a model for the role of deliberate TE reactivation in germ-line companion cells as part of a developmental mechanism for first revealing and then silencing TEs via small RNA, which may contribute to reprogramming during early development in plants and animals.