Why have some plants lost the organizational stability in plastid genomes (plastomes) that evolved in their algal ancestors? During the endosymbiotic transformation of a cyanobacterium into the eukaryotic plastid, most cyanobacterial genes were transferred to the nucleus or otherwise lost from the plastome, and the resulting plastome architecture in land plants confers organizational stability, as evidenced by the conserved gene order among bryophytes and lycophytes, whereas ferns, gymnosperms, and angiosperms share a single, 30-kb inversion. Although some additional gene losses have occurred, gene additions to angiosperm plastomes were previously unknown. Plastomes in the Campanulaceae sensu lato have incorporated dozens of large ORFs (putative proteincoding genes). These insertions apparently caused many of the 125+ large inversions now known in this small eudicot clade. This phylogenetically restricted phenomenon is not biogeographically localized, which indicates that these ORFs came from the nucleus or (less likely) a cryptic endosymbiont.foreign DNA | Cyphiaceae | Lobeliaceae | phylogeny T he extant diversity of algae and land plants chronicles the ongoing endosymbiotic transformation of a cyanobacterium into eukaryotic plastids (1), which are commonly known as chloroplasts because of their primary photosynthetic function. The early steps in plastome evolution involved the loss or transfer to the nucleus of most cyanobacterial genes, but an intron maturase (matK) and the two largest genes [ycf1 (an inner membrane translocon component); ref.2) and ycf2 (still of unknown function)] were incorporated before the origin of land plants (3). The characteristic plastome architecture of land plants (Fig.