title: Safety in numbers: multiple occurrences of highly similar homologs among Azotobacter vinelandii carbohydrate metabolism proteins probably confer adaptive benefits.
abstract: BACKGROUND
  Gene duplication and horizontal gene transfer are common processes in bacterial and archaeal genomes, and are generally assumed to result in either diversification or loss of the redundant gene copies. However, a recent analysis of the genome of the soil bacterium Azotobacter vinelandii DJ revealed an abundance of highly similar homologs among carbohydrate metabolism genes. In many cases these multiple genes did not appear to be the result of recent duplications, or to function only as a means of stimulating expression by increasing gene dosage, as the homologs were located in varying functional genetic contexts. Based on these initial findings we here report in-depth bioinformatic analyses focusing specifically on highly similar intra-genome homologs, or synologs, among carbohydrate metabolism genes, as well as an analysis of the general occurrence of very similar synologs in prokaryotes.
  
  
  RESULTS
  Approximately 900 bacterial and archaeal genomes were analysed for the occurrence of synologs, both in general and among carbohydrate metabolism genes specifically. This showed that large numbers of highly similar synologs among carbohydrate metabolism genes are very rare in bacterial and archaeal genomes, and that the A. vinelandii DJ genome contains an unusually large amount of such synologs. The majority of these synologs were found to be non-tandemly organized and localized in varying but metabolically relevant genomic contexts. The same observation was made for other genomes harbouring high levels of such synologs. It was also shown that highly similar synologs generally constitute a very small fraction of the protein-coding genes in prokaryotic genomes. The overall synolog fraction of the A. vinelandii DJ genome was well above the data set average, but not nearly as remarkable as the levels observed when only carbohydrate metabolism synologs were considered.
  
  
  CONCLUSIONS
  Large numbers of highly similar synologs are rare in bacterial and archaeal genomes, both in general and among carbohydrate metabolism genes. However, A. vinelandii and several other soil bacteria harbour large numbers of highly similar carbohydrate metabolism synologs which seem not to result from recent duplication or transfer events. These genes may confer adaptive benefits with respect to certain lifestyles and environmental factors, most likely due to increased regulatory flexibility and/or increased gene dosage.
authors.forename: Mali
authors.initials: M
authors.lastname: Mærk
authors.forename: Jostein
authors.initials: J
authors.lastname: Johansen
authors.forename: Helga
authors.initials: H
authors.lastname: Ertesvåg
authors.affiliation: Department of Cancer Research and Molecular Medicine, Norwegian University of Science and Technology, NO-7491, Trondheim, Norway. finn.drablos@ntnu.no.
authors.forename: Finn
authors.initials: F
authors.lastname: Drabløs
authors.forename: Svein
authors.initials: S
authors.lastname: Valla
mesh_terms: D000222:Adaptation, Physiological; D019843:Archaeal Proteins / Q000235:genetics; D016948:Azotobacter vinelandii / Q000235:genetics*; D001426:Bacterial Proteins / Q000235:genetics*; D050260:Carbohydrate Metabolism / Q000235:genetics*; D017124:Conserved Sequence; D016680:Genome, Bacterial; D020543:Proteome / Q000235:genetics; D011549:Pseudomonas / Q000235:genetics; D017386:Sequence Homology, Amino Acid
chemical_list: D019843:Archaeal Proteins; D001426:Bacterial Proteins; D020543:Proteome