Expression, Imprinting, and Evolution of Rice Homologs of the Polycomb Group Genes

Polycomb group proteins （PcG） play important roles in epigenetic regulation of gene expression. Some core PeG proteins, such as Enhancer of Zeste （E（z））, Suppressor of Zeste （12） （Su（z）12）, and Extra Sex Combs （ESC）, are conserved in plants. The rice genome contains two E（z）-Iike genes, OsiEZ1 and OsCLF, two homologs of Su（z）12, OsEMF2a and OsEMF2b, and two ESC-like genes, OsFIE1 and OsFIE2. OsFIE1 is expressed only in endosperm; the maternal copy is expressed while the paternal copy is not active. Other rice PcG genes are expressed in a wide range of tissues and are not imprinted in the endosperm. The two E（z）-Iike genes appear to have duplicated before the separation of the dicots and monocots; the two homologs of Su（z）12 possibly duplicated during the evolution of the Gramineae and the two ESC- like genes are likely to have duplicated in the ancestor of the grasses. No homologs of the Arabidopsis seed-expressed PcG genes MEA and FIS2 were identified in the rice genome. We have isolated T-DNA insertion lines in the rice homologs of three PcG genes. There is no autonomous endosperm development in these T-DNA insertion lines. One line with a T-DNA insertion in OsEMF2b displays pleiotropic phenotypes including altered flowering time and abnormal flower organs, suggesting important roles in rice development for this gene.

Imprinting in Plants and Its Underlying Mechanisms

Genomic imprinting（or imprinting） refers to an epigenetic phenomenon by which the allelic expression of a gene depends on the parent of origin.It has evolved independently in placental mammals and flowering plants.In plants,imprinting is mainly found in endosperm. Recent genome-wide surveys in Arabidopsis,rice,and maize identified hundreds of imprinted genes in endosperm.Since these genes are of diverse functions,endosperm development is regulated at different regulatory levels.The imprinted expression of only a few genes is conserved between Arabidopsis and monocots,suggesting that imprinting evolved quickly during speciation.In Arabidopsis,DEMETER （DME） mediates hypomethylation in the maternal genome at numerous loci（mainly transposons and repeats） in the central cell and results in many differentially methylated regions between parental genomes in the endosperm,and subsequent imprinted expression of some genes.In addition,histone modification mediated by Polycomb group（PcG） proteins is also involved in regulating imprinting.DME-induced hypomethylated alleles in the central cell are considered to produce small interfering RNAs（siRNAs） which are imported to the egg to reinforce DNA methylation.In parallel,the activity of DME in the vegetative cell of the male gametophyte demethylates many regions which overlap with the demethylated regions in the central cell.siRNAs from the demethylated regions are hypothesized to be also transferred into sperm to reinforce DNA methylation.Imprinting is partly the result of genome-wide epigenetic reprogramming in the central cell and vegetative cell and evolved under different selective pressures.