Arguably, the two most important organelles of the eukaryotic cell are not its original inhabitants. Mitochondria and chloroplasts have evolved by endo symbiosis and have a prokaryotic ancestry. This is refl ected perfectly in their genome organization and the basic functioning of their genetic system. Over the long course of evolution, they have given away most of their genes. The genes were either lost because they were no longer needed or were relocated to the nucleus. Nuclear transfer of their DNA is an ongoing process, and the two organellar genomes are still evolving. The DNA sequences are in a constant state of motion because of interorganellar and horizontal gene transfer within and between different plant species. In contrast to the chloroplast genomes, which are much conserved and do not generally exhibit any structural or functional anomalies, mitochondria are notorious for their lack of synteny, frequent DNA rearrangements, constantly varying intergenic regions and accumulation of heterologous DNA sequences. Since the nucleus cannot work in isolation, all the random and mischievous genetic activities of organellar genomes have had a direct or indirect impact on nuclear genome evolution. They act as one of the major mechanisms by which genetic novelty is brought about in the nuclear genome. As they say, to produce good music, both black and white chords have to work in harmony. Similarly, irrespective of individual nature of the three genomes in a cell, these three always work in harmony for optimum functioning of the plant cell.