The extraordinary PX-478 conservation of 16S rRNA in cyanobacteria seems to indicate that concerted evolution is a more likely explanation. To verify this suggestion we examined variation in the internal

transcribed spacer region, located between the 16S and 23S rRNA gene. Though previous studies have suggested conservation of some regions in the ITS sequence, several regions should not be affected by selection and evolve neutrally. If the entire ITS sequence showed the same degree of conservation as does the 16S gene sequence, then purifying selection —which would only act on the functional parts— could be rejected as a driving force. However, the strong conservation found in cyanobacterial 16S rRNA gene sequences could not be confirmed for the ITS-regions of four cyanobacterial

taxa (Additional file 9). For cyanobacteria and the eubacterial phyla studied here, both concerted evolution and strong purifying selection, selleck compound appear to be the main contributing factors. Although, cyanobacteria are assumed to be an ancient phylum which presumably raised oxygen levels in the atmosphere more than 2.3 billion years ago [54], variation in 16S rRNA copies is extremely low. Indeed, phylogenetic tree reconstructions GS-4997 in vivo for 16S rRNA result in relatively short estimated branch lengths within this phylum, compared to other eubacterial phyla (Figure 2). Short evolutionary distances for 16S rRNA sequences are Flavopiridol (Alvocidib) consistent with a pattern that has been found for morphological characters in cyanobacteria before. In 1994, J.W. Schopf compared the tempo and mode of evolution in cyanobacteria from the Precambrian, to evolutionary patterns observed in fossils during the Phanerozoic. The latter have been described by G.G. Simpson in his book “The tempo and mode of evolution” [55]. Schopf found that evolutionary predictions which Simpson made for metazoan fossils from the Phanerozoic, can also be applied to cyanobacteria. Morphologically, cyanobacteria seem to evolve not only at a “bradytelic”, but “hypobradytelic” mode, meaning at exceedingly low

evolutionary rates. Fossils from the Precambrian strongly resemble present morphotypes. The oldest undisputed cyanobacterial fossils date back circa 2.0 billion years [18, 19]. Morphological appearance of these microfossils already suggests the presence of at least four of the morphological sections described by Castenholz [20]. It seems that cyanobacteria reached their maximum morphological complexity two billion years ago, and many of today’s species could be described as so-called ‘living fossils’. It remains to be seen whether the low evolutionary rates as seen in 16S rRNA sequences and morphological features, is also seen at the genomic and metabolic level. This question can be further resolved as further genomic sequences become available for the cyanobacteria.