title: Local hopping mobile DNA implicated in pseudogene formation and reductive evolution in an obligate cyanobacteria-plant symbiosis.
abstract: BACKGROUND
  Insertion sequences (ISs) are approximately 1 kbp long "jumping" genes found in prokaryotes. ISs encode the protein Transposase, which facilitates the excision and reinsertion of ISs in genomes, making these sequences a type of class I ("cut-and-paste") Mobile Genetic Elements. ISs are proposed to be involved in the reductive evolution of symbiotic prokaryotes. Our previous sequencing of the genome of the cyanobacterium 'Nostoc azollae' 0708, living in a tight perpetual symbiotic association with a plant (the water fern Azolla), revealed the presence of an eroding genome, with a high number of insertion sequences (ISs) together with an unprecedented large proportion of pseudogenes. To investigate the role of ISs in the reductive evolution of 'Nostoc azollae' 0708, and potentially in the formation of pseudogenes, a bioinformatic investigation of the IS identities and positions in 47 cyanobacterial genomes was conducted. To widen the scope, the IS contents were analysed qualitatively and quantitatively in 20 other genomes representing both free-living and symbiotic bacteria.
  
  
  RESULTS
  Insertion Sequences were not randomly distributed in the bacterial genomes and were found to transpose short distances from their original location ("local hopping") and pseudogenes were enriched in the vicinity of IS elements. In general, symbiotic organisms showed higher densities of IS elements and pseudogenes than non-symbiotic bacteria. A total of 1108 distinct repeated sequences over 500 bp were identified in the 67 genomes investigated. In the genome of 'Nostoc azollae' 0708, IS elements were apparent at 970 locations (14.3%), with 428 being full-length. Morphologically complex cyanobacteria with large genomes showed higher frequencies of IS elements, irrespective of life style.
  
  
  CONCLUSIONS
  The apparent co-location of IS elements and pseudogenes found in prokaryotic genomes implies earlier IS transpositions into genes. As transpositions tend to be local rather than genome wide this likely explains the proximity between IS elements and pseudogenes. These findings suggest that ISs facilitate the reductive evolution in for instance in the symbiotic cyanobacterium 'Nostoc azollae' 0708 and in other obligate prokaryotic symbionts.
authors.affiliation: Department of Ecology, Environment and Plant Sciences, Stockholm University, Science for Life Laboratory, SE-17165, Solna, Sweden. Theoden.Vigil-Stenman@su.se.
authors.forename: Theoden
authors.initials: T
authors.lastname: Vigil-Stenman
authors.affiliation: Department of Biology and Environmental Science, Linné University, Science for Life Laboratory, SE-17165, Solna, Sweden. John.Larsson@scilifelab.se.
authors.forename: John
authors.initials: J
authors.lastname: Larsson
authors.affiliation: BILS/Swedish Museum of Natural History, Science for Life Laboratory, SE-17165, Solna, Sweden. Johan.Nylander@nrm.se.
authors.forename: Johan A A
authors.initials: JA
authors.lastname: Nylander
authors.affiliation: Department of Ecology, Environment and Plant Sciences, Stockholm University, Science for Life Laboratory, SE-17165, Solna, Sweden. Birgitta.Bergman@su.se.
authors.forename: Birgitta
authors.initials: B
authors.lastname: Bergman
mesh_terms: D000458:Cyanobacteria / Q000235:genetics* / Q000378:metabolism; D004251:DNA Transposable Elements; D019143:Evolution, Molecular; D023061:Gene Order; D059646:Genome Size; D016680:Genome, Bacterial; D016254:Mutagenesis, Insertional; D010802:Phylogeny; D011544:Pseudogenes; D012091:Repetitive Sequences, Nucleic Acid; D013559:Symbiosis / Q000235:genetics*
chemical_list: D004251:DNA Transposable Elements