Cyanobacteria are an ancient and globally distributed group of photosynthetic prokaryotes including species capable of fixing atmospheric dinitrogen (N2) into biologically available ammonia via the enzyme complex nitrogenase. The ability to form symbiotic interactions with eukaryotic hosts is a notable feature of cyanobacteria and one which, via an ancient endosymbiotic event, led to the evolution of chloroplasts and eventually to the plant dominated biosphere of the globe. Some cyanobacteria are still symbiotically competent and form symbiotic associations with eukaryotes ranging from unicellular organisms to complex plants. Among contemporary plant-cyanobacteria associations, the symbiosis formed between the small fast-growing aquatic fern Azolla and its cyanobacterial symbiont (cyanobiont), harboured in specialized cavities in each Azolla leaf, is the only one which is perpetual and in which the cyanobiont has lost its free-living capacity, suggesting a long-lasting co-evolution between the two partners. In this study, the genome of the cyanobiont in Azolla filiculoides was sequenced to completion and analysed. The results revealed that the genome is in an eroding state, evidenced by a high proportion of pseudogenes and transposable elements. Loss of function was most predominant in genetic categories related to uptake and metabolism of nutrients, response to environmental stimuli and in the DNA maintenance machinery. Conversely, function was retained in key symbiotic processes such as nitrogen-fixation and cell differentiation. A comparative analysis shows that the size of the cyanobiont genome has remained relatively stable, and that few genes have been completely eliminated, since the symbiotic establishment. Indications of genes acquired via horizontal gene transfer were discovered in thec yanobiont genome, some of which may have originated from the bacterial community in the Azolla leaf-cavities. It is concluded that the perpetual nature of the Azolla symbiosis has resulted in pronounced ongoing streamlining of the cyanobiont genome around core symbiotic functions, a process not described previously for complex cyanobacteria or for any bacterial plant symbiont. Further, the status of the genome indicates that the cyanobiont is at an early stage of adapting to its host-restricted environment and continued co-evolution with the plant may result in additional genome reductions. However, although a vertical transmission process is already established, the unusual extracellular location of the cyanobiont and the intricate nature of the symbiosis, may still impose restrictions on such a reductive process. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Submitted. Paper 3: Manuscript.</p>
| Identifer | oai:union.ndltd.org:UPSALLA1/oai:DiVA.org:su-56851 |
| Date | January 2011 |
| Creators | Larsson, John |
| Publisher | Stockholms universitet, Botaniska institutionen, Stockholm : Department of Botany, Stockholm University |
| Source Sets | DiVA Archive at Upsalla University |
| Language | English |
| Detected Language | English |
| Type | Doctoral thesis, comprehensive summary, info:eu-repo/semantics/doctoralThesis, text |
| Format | application/pdf |
| Rights | info:eu-repo/semantics/openAccess |
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