{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,1,23]],"date-time":"2026-01-23T16:56:43Z","timestamp":1769187403519,"version":"3.49.0"},"reference-count":57,"publisher":"Microbiology Society","issue":"Pt_6","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2014,6,1]]},"abstract":"<jats:p>The symbiosis <jats:italic>Azolla<\/jats:italic>\u2013<jats:italic>Anabaena azollae<\/jats:italic>, with a worldwide distribution in pantropical and temperate regions, is one of the most studied, because of its potential application as a biofertilizer, especially in rice fields, but also as an animal food and in phytoremediation. The cyanobiont is a filamentous, heterocystic cyanobacterium that inhabits the foliar cavities of the pteridophyte and the indusium on the megasporocarp (female reproductive structure). The classification and phylogeny of the cyanobiont is very controversial: from its morphology, it has been named <jats:italic>Nostoc azollae<\/jats:italic>, <jats:italic>Anabaena azollae<\/jats:italic>, <jats:italic>Anabaena variabilis<\/jats:italic> status <jats:italic>azollae<\/jats:italic> and recently <jats:italic>Trichormus azollae<\/jats:italic>, but, from its 16S rRNA gene sequence, it has been assigned to <jats:named-content xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" content-type=\"genus\" xlink:type=\"simple\">\n                     <jats:ext-link ext-link-type=\"uri\" xlink:href=\"http:\/\/dx.doi.org\/10.1601\/nm.748\" xlink:type=\"simple\">\n                        <jats:italic>Nostoc<\/jats:italic>\n                     <\/jats:ext-link>\n                  <\/jats:named-content> and\/or <jats:named-content xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" content-type=\"genus\" xlink:type=\"simple\">\n                     <jats:ext-link ext-link-type=\"uri\" xlink:href=\"http:\/\/dx.doi.org\/10.1601\/nm.729\" xlink:type=\"simple\">\n                        <jats:italic>Anabaena<\/jats:italic>\n                     <\/jats:ext-link>\n                  <\/jats:named-content>, and from its phycocyanin gene sequence, it has been assigned as non-<jats:italic>Nostoc<\/jats:italic> and non-<jats:italic>Anabaena<\/jats:italic>. The literature also points to a possible co-evolution between the cyanobiont and the <jats:italic>Azolla<\/jats:italic> host, since dendrograms and phylogenetic trees of fatty acids, short tandemly repeated repetitive (STRR) analysis and restriction fragment length polymorphism (RFLP) analysis of <jats:italic>nif<\/jats:italic> genes and the 16S rRNA gene give a two-cluster association that matches the two-section ranking of the host (<jats:italic>Azolla<\/jats:italic>). Another controversy surrounds the possible existence of more than one genus or more than one species strain. The use of freshly isolated or cultured cyanobionts is an additional problem, since their morphology and protein profiles are different. This review gives an overview of how morphological, chemical and genetic analyses influence the classification and phylogeny of the cyanobiont and future research.<\/jats:p>","DOI":"10.1099\/ijs.0.059238-0","type":"journal-article","created":{"date-parts":[[2014,4,16]],"date-time":"2014-04-16T06:06:36Z","timestamp":1397628396000},"page":"1830-1840","source":"Crossref","is-referenced-by-count":38,"title":["Classification and phylogeny of the cyanobiont Anabaena azollae Strasburger: an answered question?"],"prefix":"10.1099","volume":"64","author":[{"given":"Ana L.","family":"Pereira","sequence":"first","affiliation":[{"name":"Interdisciplinary Centre of Marine and Environmental Research (CIIMAR\/CIMAR), University of Porto, Rua dos Bragas 289, 4050-123 Porto, Portugal"}]},{"given":"Vitor","family":"Vasconcelos","sequence":"additional","affiliation":[{"name":"Department of Biology, Faculty of Sciences of the University of Porto, Rua do Campo Alegre, 4069-007 Porto, Portugal"},{"name":"Interdisciplinary Centre of Marine and Environmental Research (CIIMAR\/CIMAR), University of Porto, Rua dos Bragas 289, 4050-123 Porto, Portugal"}]}],"member":"345","reference":[{"key":"r1","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1016\/S0378-1097(03)00784-5","article-title":"The cyanobiont in an Azolla fern is neither Anabaena nor Nostoc","volume":"229","author":"Baker","year":"2003","journal-title":"FEMS Microbiol Lett"},{"key":"r2","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1007\/BF02277375","article-title":"Pigment distribution and nitrogen fixation in Anabaena azollae","volume":"61","author":"Becking","year":"1981","journal-title":"Plant Soil"},{"key":"r3","first-page":"447","article-title":"Cyanobacterial-plant symbioses: signalling and development","volume-title":"The Cyanobacteria. 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A challenge for the future","volume":"23","author":"Carrapi\u00e7o","year":"2000","journal-title":"Rev Cienc Agric"},{"key":"r6","doi-asserted-by":"crossref","first-page":"246","DOI":"10.1099\/00207713-42-2-246","article-title":"Differentiation of free-living Anabaena and Nostoc cyanobacteria on the basis of fatty acid composition","volume":"42","author":"Caudales","year":"1992","journal-title":"Int J Syst Bacteriol"},{"key":"r7","doi-asserted-by":"crossref","first-page":"1489","DOI":"10.1099\/00221287-138-7-1489","article-title":"Cellular fatty acid composition of symbiotic cyanobacteria isolated from the aquatic fern Azolla","volume":"138","author":"Caudales","year":"1992","journal-title":"J Gen Microbiol"},{"key":"r8","doi-asserted-by":"crossref","first-page":"364","DOI":"10.1099\/00207713-45-2-364","article-title":"Fatty acid composition of symbiotic cyanobacteria from different host plant (Azolla) species: evidence for coevolution of host and symbiont","volume":"45","author":"Caudales","year":"1995","journal-title":"Int J Syst Bacteriol"},{"key":"r9","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1007\/BF02812700","article-title":"Cainozoic ferns and their distribution","volume":"53","author":"Collinson","year":"2001","journal-title":"Brittonia"},{"key":"r10","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1016\/S0021-9673(99)00930-9","article-title":"Separation and identification of hydrocarbons and other volatile compounds from cultured blue-green alga Nostoc sp. by gas chromatography-mass spectrometry using serially coupled capillary columns with consecutive nonpolar and semipolar stationary phases","volume":"862","author":"Dembitsky","year":"1999","journal-title":"J Chromatogr A"},{"key":"r11","doi-asserted-by":"crossref","first-page":"1023","DOI":"10.1093\/jxb\/erm282","article-title":"Proteomic analysis of the cyanobacterium of the Azolla symbiosis: identity, adaptation, and nifH modification","volume":"59","author":"Ekman","year":"2008","journal-title":"J Exp Bot"},{"key":"r12","first-page":"377","article-title":"Anabaena variabilis status azollae","author":"Fjerdingstad","year":"1976","journal-title":"Arch Hydrobiol"},{"key":"r13","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1016\/0168-9452(85)90103-7","article-title":"The structural nif genes of four symbiotic Anabaena azollae show a highly conserved physical arrangement","volume":"39","author":"Franche","year":"1985","journal-title":"Plant Sci"},{"key":"r14","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1007\/BF00422316","article-title":"Antigenic differences between Anabaena azollae fresh from the Azolla fern leaf cavity and free-living cyanobacteria","volume":"128","author":"Gates","year":"1980","journal-title":"Arch Microbiol"},{"key":"r15","doi-asserted-by":"crossref","first-page":"2141","DOI":"10.1128\/AEM.57.8.2141-2146.1991","article-title":"Identification of a common cyanobacterial symbiont associated with Azolla spp. through molecular and morphological characterization of free-living and symbiotic cyanobacteria","volume":"57","author":"Gebhardt","year":"1991","journal-title":"Appl Environ Microbiol"},{"key":"r16","first-page":"251","article-title":"Diversity of soil Nostoc strains: phylogenetic and phenotypic variability","volume":"159","author":"Hrouzek","year":"2005","journal-title":"Arch Hydrobiol"},{"key":"r17","first-page":"5","article-title":"The modern classification of cyanoprokaryotes (cyanobacteria)","volume":"34","author":"Kom\u00e1rek","year":"2005","journal-title":"Oceanol Hydrobiol Stud"},{"key":"r18","doi-asserted-by":"crossref","first-page":"349","DOI":"10.4490\/ALGAE.2006.21.4.349","article-title":"Cyanobacterial taxonomy: current problems and prospects for the integration of traditional and molecular approaches","volume":"21","author":"Kom\u00e1rek","year":"2006","journal-title":"Algae"},{"key":"r19","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1007\/s10750-009-0030-4","article-title":"Modern taxonomy revision of planktic nostocacean cyanobacteria: a short review of genera","volume":"639","author":"Kom\u00e1rek","year":"2010a","journal-title":"Hydrobiologia"},{"key":"r20","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1007\/s10750-009-0031-3","article-title":"Recent changes (2008) in cyanobacteria taxonomy based on a combination of molecular background with phenotype and ecological consequences (genus and species concept)","volume":"639","author":"Kom\u00e1rek","year":"2010b","journal-title":"Hydrobiologia"},{"key":"r21","first-page":"247","article-title":"Modern approach to the classification system of cyanophytes. 4. 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