The mitochondrial genomes of angiosperms are the largest mitochondrial genomes so far reported and are highly variable in size among plant species. The comparative analysis of the angiosperm mitochondrial genomes at the nucleotide level has now become feasible for addressing long-standing questions, owing to the publication of five dicot and three monocot genomes. Whereas the identified genes and introns are rather well conserved, intergenic regions are highly variable in sequence, even between two close relatives. Promiscuous DNA and horizontally transferred sequence constitute part of the intergenic regions, but the origin of the majority of these regions is unknown. On the other hand, duplication and extensive rearrangement of preexisting sequences may be one of the explanations for the occurrence of unknown sequences. Functional aspects of the mitochondrial genome, such as RNA editing and expression of unique open reading frames (ORFs), can be changed under certain nuclear genotypes.