Deviations from Chargaff's second parity rule in organellar DNA: Insights into the evolution of organellar genomes

Chargaff' s second parity rule (PR2) states that complementary nucleotides are met with almost equal frequencies in single stranded DNA. This is indeed the case for all bacterial and eukaryotic genomes studied, although the genomic patterns may differ among genomes in terms of local deviations. The behaviour of organellar genomes regarding the second parity rule has not been studied in detail up to now. We tested all available organellar genomes and found that a large number of mitochondrial genomes significantly deviate from the 2nd parity rule in contrast to the eubacterial ones, although mitochondria are believed to have evolved from proteobacteria. Moreover, mitochondria may be divided into three distinct sub-groups according to their overall deviation from the aforementioned parity rule. On the other hand, chloroplast genomes share the pattern of eubacterial genomes and, interestingly, so do mitochondrial genomes originating from plants and some fungi. The deviation from the second parity is found to be weakly correlated with the overall excess of purines against pyrimidines. The behaviour of the large majority of the mitochondrial genomes may be attributed to their distinct mode of replication, which is fundamentally different from the one of the eubacteria. Differences between chloroplast and mitochondrial genomes might also be explained on the basis of different replication mechanisms and correlated to differences in the genome size and compaction. The results presented herein may provide some insight into different modes of evolution of genome structure between chloroplasts and mitochondria.