In plants, the most abundant class of small RNAs are 24-nucleotide short interfering (si)RNA. These siRNAs are produced at thousands of discrete genomic locations through the action of the plant-specific DNA-dependent RNA polymerase IV (Pol IV). Pol IV-dependent siRNAs catalyze repressive DNA methylation on transposable elements and other repetitive sequences, but might trigger diverse chromatin modifications at distinct genomic locations, such as DNA demethylation or histone modification. Pol IV-dependent siRNAs are expressed abundantly, and sometimes exclusively, in the developing endosperm, where they are produced from only the maternal chromosomes. The biological role of Pol IV-dependent siRNAs is unclear, but might involve interaction between different genomes or alleles, or stabilizing and buffering the genome from genetic and epigenetic modifications.