藍藻は土壌や岩石表面など陸域にも多く、極地から熱帯まで広く分布している ('''下図''')。変水性 (体内の水分量が大きく変動しても、つまり乾燥しても仮死状態で生存し、再び水が得られると急速に復活する性質) による高い乾燥耐性を示すものもおり、100年近く乾燥状態に置かれたものが復活したとの報告もある<ref name="Ward2000">Ward, D. M. & Castenholz, R. W. (2000) Cyanobacteria in geothermal habitats. In: ''The Ecology of Cyanobacteria'', Springer Netherlands. ISBN 0-09-941464-3 pp. 37-59.</ref>。土壌表層では藍藻が土壌クラスト (soil crust) を形成し ('''下図''')、土壌の安定化や窒素栄養分の供給を行うため、特に砂漠や荒野での[[植生遷移]]の初期段階において重要な働きを果たすことがある<ref name="Whitton2000" /><ref name="Whitton2012" />。藍藻の中には、岩石内生 (岩の中に生育) のものもおり、砂漠や極地からも見つかっている。
藍藻は土壌や岩石表面など陸域にも多く、極地から熱帯まで広く分布している ('''下図''')。変水性 (体内の水分量が大きく変動しても、つまり乾燥しても仮死状態で生存し、再び水が得られると急速に復活する性質) による高い乾燥耐性を示すものもおり、100年近く乾燥状態に置かれたものが復活したとの報告もある<ref name="Ward2000">Ward, D. M. & Castenholz, R. W. (2000) Cyanobacteria in geothermal habitats. In: ''The Ecology of Cyanobacteria'', Springer Netherlands. ISBN 0-09-941464-3 pp. 37-59.</ref>。土壌表層では藍藻が土壌クラスト (soil crust) を形成し ('''下図''')、土壌の安定化や窒素栄養分の供給を行うため、特に砂漠や荒野での[[植生遷移]]の初期段階において重要な働きを果たすことがある<ref name="Whitton2000" /><ref name="Whitton2012" />。藍藻の中には、岩石内生 (岩の中に生育) のものもおり、砂漠や極地からも見つかっている<ref name="Wierzchos2006">{{cite journal|author=Wierzchos, J., Ascaso, C. & McKay, C. P.|year=2006|title=Endolithic cyanobacteria in halite rocks from the hyperarid core of the Atacama Desert|journal=Astrobiology|volume=6|pages=415-422|doi=10.1089/ast.2006.6.415}}</ref>。
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| caption2 = [[ソテツ属]]のサンゴ状根 (内部に藍藻が共生).
| caption2 = [[ソテツ属]]のサンゴ状根 (内部に藍藻が共生).
}}
}}
<div id="symbiosis with phototroph">光合成生物に藍藻が共生している例も知られている。このような共生では、'''藍藻が窒素固定によって生成した窒素化合物を宿主に与えている'''<ref name="Adams2012" /><ref name="Rai2000">{{cite journal|author=Rai, A. N., Söderbäck, E. & Bergman, B.|year=2000|title=Cyanobacterium-plant symbioses|journal=New Phytologist|volume=147|pages=449-481|doi=10.1046/j.1469-8137.2000.00720.x}}</ref>。藍藻が共生している[[陸上植物]]として、[[ウスバゼニゴケ科]]<ref name="Adams2008">{{cite journal|author=Adams, D. G. & Duggan, P. S.|year=2008|title=Cyanobacteria-bryophyte symbioses|journal=J. Exp. Bot.|volume=59|pages=1047-1058|doi=10.1093/jxb/ern005}}</ref> ([[苔類]])、[[ツノゴケ類]]<ref name="Adams2008" />、[[アカウキクサ属]]<ref name="Peters1991">{{cite journal|author=Peters, G.A.|year=1991|title=''Azolla'' and other plant-cyanobacteria symbioses - aspects of form and function|journal=Plant Soil|volume=137|pages=25-36|doi=10.1007/BF02187428}}</ref><ref name="Papaefthimiou2008">{{cite journal|author=Papaefthimiou, D., Van Hove, C., Lejeune, A., Rasmussen, U. & Wilmotte, A.|year=2008|title=Diversity and host specificity of genus ''Azolla'' cyanobionts|journal=J. Phycol.|volume=44|pages=60-70|doi=10.1111/j.1529-8817.2007.00448.x}}</ref> ([[シダ綱|薄曩シダ]]) ('''右図''')、[[ソテツ類]]<ref name="Costa2003">{{cite book|author=Costa, J.-L. & Lindblad, P.|year=2003|chapter=Cyanobacteria in symbiosis with cycads|editor=Rai, A.N., Bergman, B. & Rasmussen, U.|title=Cyanobacteria in Symbiosis|publisher=Kluwer Academic Publishers, Dordrecht|isbn=1-4020-0777-9|pages=195-205}}</ref> ('''右図''')、[[グンネラ]]<ref name="Bergman2002">{{cite book|author=Bergman, B.|year=2002|chapter=The ''Nostoc''-''Gunnera'' symbiosis|editor=Rai, A.N., Bergman, B. & Rasmussen, U.|title=Cyanobacteria in Symbiosis|publisher=Kluwer Academic Publishers|isbn=1-4020-0777-9|pages=207-232}}</ref><ref name="Bergman2008">{{cite journal|author=Bergman, B. & Osborne, B.|year=2002|title=The ''Gunnera''-''Nostoc'' symbiosis|journal=Biol. Environ. Proc. R Ir Acad.|volume=102B|pages=35-39|url=https://www.jstor.org/stable/20500139}}</ref> ([[被子植物]]) などが知られている。これらの中には、藍藻の感染を促進するために植物が藍藻の[[#生殖|連鎖体]]や[[線毛]]形成を誘導する例が知れられている<ref name="Duggan2007" /><ref name="Adams2012" />。またアカウキクサ類の共生藍藻は[[宿主]]体外では生存不可な絶対共生性であり、宿主と[[共進化]]していることが知られている<ref name="Adams2012" /><ref name="Papaefthimiou2008" />。ソテツ類はいくつかの[[毒素]]をもつことが知られているが、このうち BMAA (β-methylamino-L-alanine) はソテツ自身が生成したものではなく、共生藍藻が生成したものであると考えられている<ref name="Cox2003">{{cite journal|author=Cox, P.A., Banack, S.A. & Murch, S.J.|year=2003|title=Biomagnification of cyanobacterial neurotoxins and neurodegenerative disease among the Chamorro people of Guam|journal=Proc. Natl. Acad. Sci. U.S.A.|volume=100|pages=13380-13383|doi=10.1073/pnas.2235808100}}</ref>。</div>
<div id="symbiosis with phototroph">光合成生物に藍藻が共生している例も知られている。このような共生では、'''藍藻が窒素固定によって生成した窒素化合物を宿主に供給'''している<ref name="Adams2012" /><ref name="Rai2000">{{cite journal|author=Rai, A. N., Söderbäck, E. & Bergman, B.|year=2000|title=Cyanobacterium-plant symbioses|journal=New Phytologist|volume=147|pages=449-481|doi=10.1046/j.1469-8137.2000.00720.x}}</ref>。藍藻が共生している[[陸上植物]]として、[[ウスバゼニゴケ科]]<ref name="Adams2008">{{cite journal|author=Adams, D. G. & Duggan, P. S.|year=2008|title=Cyanobacteria-bryophyte symbioses|journal=J. Exp. Bot.|volume=59|pages=1047-1058|doi=10.1093/jxb/ern005}}</ref> ([[苔類]])、[[ツノゴケ類]]<ref name="Adams2008" />、[[アカウキクサ属]]<ref name="Peters1991">{{cite journal|author=Peters, G.A.|year=1991|title=''Azolla'' and other plant-cyanobacteria symbioses - aspects of form and function|journal=Plant Soil|volume=137|pages=25-36|doi=10.1007/BF02187428}}</ref><ref name="Papaefthimiou2008">{{cite journal|author=Papaefthimiou, D., Van Hove, C., Lejeune, A., Rasmussen, U. & Wilmotte, A.|year=2008|title=Diversity and host specificity of genus ''Azolla'' cyanobionts|journal=J. Phycol.|volume=44|pages=60-70|doi=10.1111/j.1529-8817.2007.00448.x}}</ref> ([[シダ綱|薄曩シダ]]) ('''右図''')、[[ソテツ類]]<ref name="Costa2003">{{cite book|author=Costa, J.-L. & Lindblad, P.|year=2003|chapter=Cyanobacteria in symbiosis with cycads|editor=Rai, A.N., Bergman, B. & Rasmussen, U.|title=Cyanobacteria in Symbiosis|publisher=Kluwer Academic Publishers, Dordrecht|isbn=1-4020-0777-9|pages=195-205}}</ref> ('''右図''')、[[グンネラ]]<ref name="Bergman2002">{{cite book|author=Bergman, B.|year=2002|chapter=The ''Nostoc''-''Gunnera'' symbiosis|editor=Rai, A.N., Bergman, B. & Rasmussen, U.|title=Cyanobacteria in Symbiosis|publisher=Kluwer Academic Publishers|isbn=1-4020-0777-9|pages=207-232}}</ref><ref name="Bergman2008">{{cite journal|author=Bergman, B. & Osborne, B.|year=2002|title=The ''Gunnera''-''Nostoc'' symbiosis|journal=Biol. Environ. Proc. R Ir Acad.|volume=102B|pages=35-39|url=https://www.jstor.org/stable/20500139}}</ref> ([[被子植物]]) などが知られている。これらの中には、藍藻の感染を促進するために植物が藍藻の[[#生殖|連鎖体]]や[[線毛]]形成を誘導する例が知れられている<ref name="Duggan2007" /><ref name="Adams2012" />。またアカウキクサ類の共生藍藻は[[宿主]]体外では生存不可な絶対共生性であり、宿主と[[共進化]]していることが知られている<ref name="Adams2012" /><ref name="Papaefthimiou2008" />。ソテツ類はいくつかの[[毒素]]をもつことが知られているが、このうち BMAA (β-methylamino-L-alanine) はソテツ自身が生成したものではなく、共生藍藻が生成したものであると考えられている<ref name="Cox2003">{{cite journal|author=Cox, P.A., Banack, S.A. & Murch, S.J.|year=2003|title=Biomagnification of cyanobacterial neurotoxins and neurodegenerative disease among the Chamorro people of Guam|journal=Proc. Natl. Acad. Sci. U.S.A.|volume=100|pages=13380-13383|doi=10.1073/pnas.2235808100}}</ref>。</div>
水界でも、[[珪藻]]<ref name="Jahson1995">{{cite journal|author=Jahson, S., Rai, A. N. & Bergman, B.|year=1995|title=Intracellular cyanobiont ''Richelia intracellularis'': ultrastructure and immuno-localisation of phycoerythrin, nitrogenase, Rubisco and glutamine synthetase|journal=Marine Biology|volume=124|pages=1-8|doi=10.1007/BF00349140}}</ref><ref name="Carpenter2002" /><ref name="Foster2006c">{{cite journal|author=Foster, R. A. & Zehr, J. P.|year=2006|title=Characterization of diatom-cyanobacteria symbioses on the basis of ''nifH'', ''hetR'' and 16S rRNA sequences|journal=Environ. Microbiol.|volume=8|pages=1913-1925|doi=10.1111/j.1462-2920.2006.01068.x}}</ref><ref name="Foster2011">{{cite journal|author=Foster, R.A., Kuypers, M.M.M., Vagner, T., Paerl, R.W., Muzat, N. & Zehr, J.P.|year=2011|title=Nitrogen fixation and transfer in open ocean diatom-cyanobacterial symbioses|journal=ISME J.|volume=5|pages=1484-1493|doi=10.1038/ismej.2011.26}}</ref><ref name="Foster2009">{{cite journal|author=Foster, R.A., Subramaniam, A. & Zehr, J.P.|year=2009|title=Distribution and activity of diazotrophs in the Eastern Equatorial Atlantic|journal=Environ. Microbiol.|volume=11|pages=741-750|doi=10.1111/j.1462-2920.2008.01796.x}}</ref><ref name="White2007">{{cite journal|author=White, A.E., Prahl, F.G., Letelier, R.M. & Popp, B.N.|year=2007|title=Summer surface waters in the Gulf of California: prime habitat for biological nitrogen fixation|journal=Glob. Biogeochem. Cycles|volume=21|pages=GB2017|doi=10.1029/2006GB002779}}</ref>や[[ハプト藻]]<ref name="Hagino2013">{{cite journal|author=Hagino, K., Onuma, R., Kawachi, M. & Horiguchi, T.|year=2013|title=Discovery of an endosymbiotic nitrogen-fixing cyanobacterium UCYN-A in ''Braarudosphaera bigelowii'' (Prymnesiophyceae)|journal=PLoS One|volume=8|pages=e81749|doi=10.1371/journal.pone.0081749}}</ref><ref name="Thompson2014">{{cite journal|author=Thompson, A., Carter, B. J., Turk‐Kubo, K., Malfatti, F., Azam, F. & Zehr, J. P.|year=2014|title=Genetic diversity of the unicellular nitrogen‐fixing cyanobacteria UCYN‐A and its prymnesiophyte host|journal=Environmental Microbiology|volume=16|pages=3238-3249|doi=10.1111/1462-2920.12490}}</ref>など光合成を行う[[藻類]]に藍藻が共生している例が知られている。ハフケイソウ科の珪藻に細胞内共生している藍藻は、既に自立能・光合成能を失い、'''楕円体''' (spheroid body) とよばれる細胞小器官になっている<ref name="Kneip2008">{{cite journal|author=Kneip, C., Voß, C., Lockhart, P. J. & Maier, U. G.|year=2008|title=The cyanobacterial endosymbiont of the unicellular algae Rhopalodia gibba shows reductive genome evolution|journal=BMC Evol. Biol.|volume=8|pages=30|doi=10.1111/1462-2920.12490}}</ref>。{{Snamei||Braarudosphaera}} (ハプト藻) に共生する藍藻 (UCYN-A) も光合成能を含むいくつかの機能を欠いており、おそらく宿主に大きく依存している<ref name="Zehr2008" />。
水界でも、[[珪藻]]<ref name="Jahson1995">{{cite journal|author=Jahson, S., Rai, A. N. & Bergman, B.|year=1995|title=Intracellular cyanobiont ''Richelia intracellularis'': ultrastructure and immuno-localisation of phycoerythrin, nitrogenase, Rubisco and glutamine synthetase|journal=Marine Biology|volume=124|pages=1-8|doi=10.1007/BF00349140}}</ref><ref name="Carpenter2002" /><ref name="Foster2006c">{{cite journal|author=Foster, R. A. & Zehr, J. P.|year=2006|title=Characterization of diatom-cyanobacteria symbioses on the basis of ''nifH'', ''hetR'' and 16S rRNA sequences|journal=Environ. Microbiol.|volume=8|pages=1913-1925|doi=10.1111/j.1462-2920.2006.01068.x}}</ref><ref name="Foster2011">{{cite journal|author=Foster, R.A., Kuypers, M.M.M., Vagner, T., Paerl, R.W., Muzat, N. & Zehr, J.P.|year=2011|title=Nitrogen fixation and transfer in open ocean diatom-cyanobacterial symbioses|journal=ISME J.|volume=5|pages=1484-1493|doi=10.1038/ismej.2011.26}}</ref><ref name="Foster2009">{{cite journal|author=Foster, R.A., Subramaniam, A. & Zehr, J.P.|year=2009|title=Distribution and activity of diazotrophs in the Eastern Equatorial Atlantic|journal=Environ. Microbiol.|volume=11|pages=741-750|doi=10.1111/j.1462-2920.2008.01796.x}}</ref><ref name="White2007">{{cite journal|author=White, A.E., Prahl, F.G., Letelier, R.M. & Popp, B.N.|year=2007|title=Summer surface waters in the Gulf of California: prime habitat for biological nitrogen fixation|journal=Glob. Biogeochem. Cycles|volume=21|pages=GB2017|doi=10.1029/2006GB002779}}</ref>や[[ハプト藻]]<ref name="Hagino2013">{{cite journal|author=Hagino, K., Onuma, R., Kawachi, M. & Horiguchi, T.|year=2013|title=Discovery of an endosymbiotic nitrogen-fixing cyanobacterium UCYN-A in ''Braarudosphaera bigelowii'' (Prymnesiophyceae)|journal=PLoS One|volume=8|pages=e81749|doi=10.1371/journal.pone.0081749}}</ref><ref name="Thompson2014">{{cite journal|author=Thompson, A., Carter, B. J., Turk‐Kubo, K., Malfatti, F., Azam, F. & Zehr, J. P.|year=2014|title=Genetic diversity of the unicellular nitrogen‐fixing cyanobacteria UCYN‐A and its prymnesiophyte host|journal=Environmental Microbiology|volume=16|pages=3238-3249|doi=10.1111/1462-2920.12490}}</ref>など光合成を行う[[藻類]]に藍藻が共生している例が知られている。ハフケイソウ科の珪藻に細胞内共生している藍藻は、既に自立能・光合成能を失い、'''楕円体''' (spheroid body) とよばれる細胞小器官になっている<ref name="Kneip2008">{{cite journal|author=Kneip, C., Voß, C., Lockhart, P. J. & Maier, U. G.|year=2008|title=The cyanobacterial endosymbiont of the unicellular algae Rhopalodia gibba shows reductive genome evolution|journal=BMC Evol. Biol.|volume=8|pages=30|doi=10.1111/1462-2920.12490}}</ref>。{{Snamei||Braarudosphaera}} (ハプト藻) に共生する藍藻 (UCYN-A) も光合成能を含むいくつかの機能を欠いており、おそらく宿主に大きく依存している<ref name="Zehr2008" />。
[[ファイル:Stromatolitic metadolostone (Kona Dolomite, Paleoproterozoic, 2.2 to 2.3 Ga; Lindberg Quarry, Marquette, Upper Peninsula of Michigan, USA) (8363769094).jpg|200px|thumb|right|22–23億年前のストロマトライト (米国ミシガン州).]]
[[ファイル:Stromatolitic metadolostone (Kona Dolomite, Paleoproterozoic, 2.2 to 2.3 Ga; Lindberg Quarry, Marquette, Upper Peninsula of Michigan, USA) (8363769094).jpg|200px|thumb|right|22–23億年前のストロマトライト (米国ミシガン州).]]
藍藻は、生物の進化において初めて (そして唯1回) 酸素発生型光合成能を獲得した生物群であると考えられている。藍藻と考えられる化石は約23〜27億年前の[[ストロマトライト]]化石に遡る<ref name="Yamamoto2013">{{cite journal|author=山本 純之 & 磯崎 行雄|year=2013|title=ストロマトライト研究の歴史と今後の展望|journal=地学雑誌|volume=122|pages=791-806|doi=10.5026/jgeography.122.791}}</ref><ref name="Lepot2008">{{cite journal|author=Lepot, K., Benzerara, K., Brown, G. E. & Philippot, P.|year=2008|title=Microbially influenced formation of 2,724-million-year-old stromatolites|journal=Nature Geoscience|volume=1|pages=118-121|doi=10.1038/ngeo107}}</ref> ('''右図''')。ストロマトライト様の化石はこれ以前 (〜35億年前) からも見つかるが<ref name="Schopf2006">{{cite journal|author=Schopf, J. W.|year=2006|title=Fossil evidence of Archaean life|journal=Phil. Trans. R. Soc. B|volume=361|pages=869-885|doi=10.1098/rstb.2006.1834}}</ref>、現在ではこれは非生物起源であると考えられている<ref name="Yamamoto2013" />。初期の藍藻が生成した[[酸素]]は、当初は海水中の鉄などを酸化し (その結果大規模な[[縞状鉄鉱床]]が形成され、現在利用される[[鉄鉱石]]の大部分はこれに由来する)、その後、海や大気中に放出されて地球が急速に酸化的環境に変化していった。この急速な変化は、'''大酸化事変''' (Great Oxygenation Event, GOE) とよばれる<ref name="Holland2006">{{cite journal|author=Holland, H. D.|year=2006|title=The oxygenation of the atmosphere and oceans|journal=Philosophical Transactions of the Royal Society: Biological Sciences|volume=361|pages=903-915|doi=10.1098/rstb.2006.1838}}</ref>。
藍藻は、生物の進化において初めて (そして唯1回) 酸素発生型光合成能を獲得した生物群であると考えられている。藍藻と考えられる化石は約23〜27億年前の[[ストロマトライト]]化石に遡る<ref name="Yamamoto2013">{{cite journal|author=山本 純之 & 磯崎 行雄|year=2013|title=ストロマトライト研究の歴史と今後の展望|journal=地学雑誌|volume=122|pages=791-806|doi=10.5026/jgeography.122.791}}</ref><ref name="Lepot2008">{{cite journal|author=Lepot, K., Benzerara, K., Brown, G. E. & Philippot, P.|year=2008|title=Microbially influenced formation of 2,724-million-year-old stromatolites|journal=Nature Geoscience|volume=1|pages=118-121|doi=10.1038/ngeo107}}</ref> ('''右図''')。ストロマトライト様の化石はこれ以前 (〜35億年前) からも見つかるが<ref name="Schopf2006">{{cite journal|author=Schopf, J. W.|year=2006|title=Fossil evidence of Archaean life|journal=Phil. Trans. R. Soc. B|volume=361|pages=869-885|doi=10.1098/rstb.2006.1834}}</ref>、現在ではこれは非生物起源であると考えられている<ref name="Yamamoto2013" />。初期の藍藻が生成した[[酸素]]は、当初は海水中の鉄などを酸化し (その結果大規模な[[縞状鉄鉱床]]が形成され、現在利用される[[鉄鉱石]]の大部分はこれに由来する)、その後、海や大気中に放出されて地球が急速に酸化的環境に変化していった。この急速な変化は、'''大酸化事変''' (大酸化イベント、Great Oxygenation Event, GOE) とよばれる<ref name="Holland2006">{{cite journal|author=Holland, H. D.|year=2006|title=The oxygenation of the atmosphere and oceans|journal=Philosophical Transactions of the Royal Society: Biological Sciences|volume=361|pages=903-915|doi=10.1098/rstb.2006.1838}}</ref>。
藍藻の誕生によって地球環境は激変し (好気的環境、有機物安定的供給など)、現在の地球[[生態系]]の基礎が築かれた<ref name="Inouye2006" /><ref name="Tomitani2006">{{cite journal|author=Tomitani, A., Knoll, A. H., Cavanaugh, C. M. & Ohno, T.|year=2006|title=The evolutionary diversification of cyanobacteria: molecular-phylogenetic and paleontological perspectives|journal=Proc. Natl. Acad. Sci. U.S.A.|volume=103|pages=5442-5447|doi=10.1073/pnas.0600999103}}</ref>。酸素発生型光合成生物は藍藻だけである時代が長く続いたが、その後 (ある研究では15億年以上前)、ある[[真核生物]]にある藍藻が[[細胞内共生説|細胞内共生]]し、やがてこの共生藍藻の増殖や代謝が[[宿主]]である真核生物に制御されるようになり、最終的に[[葉緑体]] ([[色素体]]) とよばれる[[細胞小器官]]へと変化した<ref name="Inouye2006" /><ref name="Archibald2009" /><ref>{{cite journal|author=Yoon, H. S., Hackett, J. D., Ciniglia, C., Pinto, G. & Bhattacharya, D.|year=2004|title=A molecular timeline for the origin of photosynthetic eukaryotes|journal=Molecular Biology and Evolution|volume=21|pages=809-818|doi=10.1093/molbev/msh075}}</ref>。この際、藍藻の[[細胞膜]]と[[外膜]]が色素体の2枚の膜になったと考えられている<ref>{{cite journal|author=Gould, S.B., Waller, R.F. & McFadden, G.I.|year=2008|title=Plastid evolution|journal=Annu. Rev. Plant Biol.|volume=59|pages=491-517|doi=10.1146/annurev.arplant.59.032607.092915}}</ref>。この現象は'''一次共生''' (primary endosymbiosis) とよばれ、これによって真核生物が酸素発生型光合成能を獲得した。生物の歴史の中で一次共生は唯1回の現象であったと考えられており、全ての葉緑体は単一の一次共生に由来する (その後、二次共生を経たものもある)。多数の遺伝子を用いた系統解析から、一次共生において共生者となった藍藻は、グロエオマルガリータ属 ({{Snamei||Gloeomargarita}}) という淡水産単細胞性藍藻に近縁な藍藻であったことが示唆されている<ref name="Ponce-Toledo2017">{{cite journal|author=Ponce-Toledo, R. I., Deschamps, P., López-García, P., Zivanovic, Y., Benzerara, K. & Moreira, D.|year=2017|title=An early-branching freshwater cyanobacterium at the origin of plastids|journal=Current Biology|volume=27|pages=386-391|doi=10.1016/j.cub.2016.11.056}}</ref>。
藍藻の誕生によって地球環境は激変し (好気的環境、有機物安定的供給、[[オゾン層]]形成など)、現在の地球[[生態系]]の基礎が築かれた<ref name="Inouye2006" /><ref name="Tomitani2006">{{cite journal|author=Tomitani, A., Knoll, A. H., Cavanaugh, C. M. & Ohno, T.|year=2006|title=The evolutionary diversification of cyanobacteria: molecular-phylogenetic and paleontological perspectives|journal=Proc. Natl. Acad. Sci. U.S.A.|volume=103|pages=5442-5447|doi=10.1073/pnas.0600999103}}</ref><ref>{{cite journal|author=Kasting, J. F.|year=1987|title=Theoretical constraints on oxygen and carbon dioxide concentrations in the Precambrian atmosphere|journal=Precambrian Research|volume=34|pages=205-229|doi=10.1016/0301-9268(87)90001-5}}</ref>。酸素発生型光合成生物は藍藻だけである時代が長く続いたが、その後 (ある研究では15億年以上前)、ある[[真核生物]]にある藍藻が[[細胞内共生説|細胞内共生]]し、やがてこの共生藍藻の増殖や代謝が[[宿主]]である真核生物に制御されるようになり、最終的に[[葉緑体]] ([[色素体]]) とよばれる[[細胞小器官]]へと変化した<ref name="Inouye2006" /><ref name="Archibald2009" /><ref>{{cite journal|author=Yoon, H. S., Hackett, J. D., Ciniglia, C., Pinto, G. & Bhattacharya, D.|year=2004|title=A molecular timeline for the origin of photosynthetic eukaryotes|journal=Molecular Biology and Evolution|volume=21|pages=809-818|doi=10.1093/molbev/msh075}}</ref>。この際、藍藻の[[細胞膜]]と[[外膜]]が色素体の2枚の膜になったと考えられている<ref>{{cite journal|author=Gould, S.B., Waller, R.F. & McFadden, G.I.|year=2008|title=Plastid evolution|journal=Annu. Rev. Plant Biol.|volume=59|pages=491-517|doi=10.1146/annurev.arplant.59.032607.092915}}</ref>。この現象は'''一次共生''' (primary endosymbiosis) とよばれ、これによって真核生物が酸素発生型光合成能を獲得した。生物の歴史の中で一次共生は唯1回の現象であったと考えられており、全ての葉緑体は単一の一次共生に由来する (その後、二次共生を経たものもある)。多数の遺伝子を用いた系統解析から、一次共生において共生者となった藍藻は、グロエオマルガリータ属 ({{Snamei||Gloeomargarita}}) という淡水産単細胞性藍藻に近縁な藍藻であったことが示唆されている<ref name="Ponce-Toledo2017">{{cite journal|author=Ponce-Toledo, R. I., Deschamps, P., López-García, P., Zivanovic, Y., Benzerara, K. & Moreira, D.|year=2017|title=An early-branching freshwater cyanobacterium at the origin of plastids|journal=Current Biology|volume=27|pages=386-391|doi=10.1016/j.cub.2016.11.056}}</ref>。
==分類==
==分類==
古くは、藍藻は最も原始的な"植物"と考えられ、藍色植物門 ({{Sname||Cyanophyta}})、藍藻綱 ({{Sname||Cyanophyceae}}) に分類されていた<ref name="Pascher1931">{{cite journal|author=Pascher, A.|year=1931|title=Systematische Übersicht über die mit Flagellaten in Zusammenhang stehenden Algenreihen und Versuch einer Einreihung dieser Algenstämme in die Stämme des Pflanzenreiches|journal=Beihefte Bot Centralbl.|volume=48|pages=317-332|doi=}}</ref><ref>{{cite book|author=Round, F.E.|year=1973|title=The Biology of the Algae. 2nd Edition|journal=Edward Arnold Publishers|pages=278|isbn=}}</ref>。しかし葉緑体の共生説が一般的となり、藍藻と他の"植物"の直接的な類縁性は認められなくなった (ただし上記のように、細胞内共生・葉緑体を通してつながっている)。これらの分類群名は植物命名規約 (現 [[国際藻類・菌類・植物命名規約]]) に基づくものであり、原核生物である藍藻に対しては近年ではほとんど用いられない<ref group="注">ただし2019年現在、原核生物の分類体系では、藍藻を分類する一般的な綱レベルの分類群名がないため、藍藻綱 ({{Sname||Cyanophyceae}}) が暫定的に用いられることがある (例:, [https://www.algaebase.org/browse/taxonomy/?id=4351 AlgaeBase])。</ref>。また古くは、粘藻綱 ({{Sname||Myxophyceae}}) や分裂藻綱 ({{Sname||Schizophyceae}}) という分類群名が使われていたこともある<ref name="Pringsheim1949">{{cite journal|author=Pringsheim, E.G.|year=1949|title=The relationship between bacteria and Myxophyceae|journal=Bacteriological Reviews|volume=13|pages=47-98|doi=}}</ref>。
古くは、藍藻は最も原始的な"植物"と考えられ、藍色植物門 ({{Sname||Cyanophyta}})、藍藻綱 ({{Sname||Cyanophyceae}}) に分類されていた<ref name="Pascher1931">{{cite journal|author=Pascher, A.|year=1931|title=Systematische Übersicht über die mit Flagellaten in Zusammenhang stehenden Algenreihen und Versuch einer Einreihung dieser Algenstämme in die Stämme des Pflanzenreiches|journal=Beihefte Bot Centralbl.|volume=48|pages=317-332|doi=}}</ref><ref>{{cite book|author=Round, F.E.|year=1973|title=The Biology of the Algae. 2nd Edition|journal=Edward Arnold Publishers|pages=278|isbn=}}</ref>。しかし葉緑体の共生説が一般的となり、藍藻と他の"植物"の直接的な類縁性は認められなくなった (ただし上記のように、細胞内共生・葉緑体を通してつながっている)。これらの分類群名は植物命名規約 (現 [[国際藻類・菌類・植物命名規約]]) に基づくものであり、原核生物である藍藻に対しては近年ではほとんど用いられない<ref group="注">ただし2019年現在、原核生物の分類体系では、藍藻を分類する一般的な綱レベルの分類群名がないため、藍藻綱 ({{Sname||Cyanophyceae}}) が暫定的に用いられることがある (例:[https://www.algaebase.org/browse/taxonomy/?id=4351 AlgaeBase])。</ref>。また古くは、粘藻綱 ({{Sname||Myxophyceae}}) や分裂藻綱 ({{Sname||Schizophyceae}}) という分類群名が使われていたこともある<ref name="Pringsheim1949">{{cite journal|author=Pringsheim, E.G.|year=1949|title=The relationship between bacteria and Myxophyceae|journal=Bacteriological Reviews|volume=13|pages=47-98|doi=}}</ref>。
藍藻は[[原核生物]]であり、[[細菌]] (バクテリア、真正細菌) [[ドメイン (分類学)|ドメイン]]に属する。細菌の中では、藍藻は比較的独立した系統群を形成しており、'''シアノバクテリア門''' ('''藍色細菌門''')(学名:{{Sname||Cyanobacteria}}) として扱われる。[[メタゲノミクス|メタゲノム研究]]によって見つかった、藍藻に近縁な[[従属栄養生物|従属栄養性]]細菌群である"[[メライナバクテリア]]綱" ({{Sname||Melainabacteria}}) や"セリキトクロマチア綱" ({{Sname||Sericytochromatia}}) などは、シアノバクテリア門に含めて扱われることがあるが、その場合は光合成能をもつ藍藻はオキシフォトバクテリア綱 ({{Sname||Oxyphotobacteria}})<ref group="注" name="Oxyphotobacteria" /> にまとめられ、これらの非光合成細菌群と併置される<ref name="Soo2014" /><ref name="Soo2017" />。ただし、シアノバクテリア門を光合成性のグループのみに限る考えもある<ref>{{cite journal|author=Garcia‐Pichel, F., Zehr, J. P., Bhattacharya, D. & Pakrasi, H. B.|year=2019|title=What's in a name? The case of cyanobacteria|journal=Journal of Phycology|volume=|pages=|doi=10.1111/jpy.12934}}</ref>。
藍藻は[[原核生物]]であり、[[細菌]] (バクテリア、真正細菌) [[ドメイン (分類学)|ドメイン]]に属する。細菌の中では、藍藻は比較的独立した系統群を形成しており、'''シアノバクテリア門''' ('''藍色細菌門''')(学名:{{Sname||Cyanobacteria}}) として扱われる。[[メタゲノミクス|メタゲノム研究]]によって見つかった、藍藻に近縁な[[従属栄養生物|従属栄養性]]細菌群である"[[メライナバクテリア]]綱" ({{Sname||Melainabacteria}}) や"セリキトクロマチア綱" ({{Sname||Sericytochromatia}}) などは、シアノバクテリア門に含めて扱われることがあるが、その場合は光合成能をもつ藍藻はオキシフォトバクテリア綱 ({{Sname||Oxyphotobacteria}})<ref group="注" name="Oxyphotobacteria" /> にまとめられ、これらの非光合成細菌群と併置される<ref name="Soo2014" /><ref name="Soo2017" />。ただし、シアノバクテリア門を光合成性のグループのみに限る考えもある<ref>{{cite journal|author=Garcia‐Pichel, F., Zehr, J. P., Bhattacharya, D. & Pakrasi, H. B.|year=2019|title=What's in a name? The case of cyanobacteria|journal=Journal of Phycology|volume=|pages=|doi=10.1111/jpy.12934}}</ref>。
ほとんどの藍藻は、クロロフィルa をもつ。また一部の藍藻は、クロロフィル a に加えて、クロロフィル b、d、または f をもつ[62][63][64]。クロロフィル d や f は生物の中で一部の藍藻のみがもつ色素であり、人間の目には見えない近赤外光を光合成に利用できる。クロロフィル b (または類似色素) をもつ藍藻は、原核緑藻ともよばれる。原核緑藻のプロクロロコックス属 (Prochlorococcus) はクロロフィル a の代わりにジビニルクロロフィル a をもつ点で特異な存在であり、光合成の反応中心でジビニルクロロフィル a を用いる唯一の生物である[65][66]。またアカリオクロリス属 (Acaryochloris) はクロロフィル a 量が少なく、反応中心でクロロフィル d を用いている[67]。
^これらの藍藻はいくつかの属に分けることが提唱されている (Walter, J. M., Coutinho, F. H., Dutilh, B. E., Swings, J., Thompson, F. L., & Thompson, C. C. (2017). “Ecogenomics and taxonomy of Cyanobacteria phylum”. Frontiers in Microbiology8: 2132.)。ただし2019年現在、これらの新しい属名は一般的ではない。
^ abcdefghiKomárek, J. (2003). “Coccoid and colonial cyanobacteria”. In Wehr, J.D. & Sheath, R.G.. Freshwater Algae of North America. Ecology and Classification. Boston, MA: Academic Press. pp. 59-116. ISBN978-0127415505
^ abcdKomárek, J., Kling, H. & Komárková, J. (2003). “Filamentous cyanobacteria”. In Wehr, J.D. & Sheath, R.G.. Freshwater Algae of North America. Ecology and Classification. Boston, MA: Academic Press. pp. 117-196. ISBN978-0127415505
^Mahasneh, I.A., Grainger, S.L.J. & Whitton, B.A. (1990). “Influence of salinity on hair formation and phosphatase-activities of the blue-green-alga (cyanobacterium) Calothrix viguieri D253”. Br. Phycol. J.25: 25-32. doi:10.1080/00071619000650021.
^ abcdFlores, E. & Herrero, A. (2010). “Compartmentalized function through cell differentiation in filamentous cyanobacteria”. Nature Reviews Microbiology8: 39-50. doi:10.1038/nrmicro2242.
^Singh, S.P. & Montgomery, B.L. (2011). “Determining cell shape: adaptive regulation of cyanobacterial cellular differentiation and morphology”. Trends in Microbiology19: 278-285. doi:10.1016/j.tim.2011.03.001.
^ abcHoiczyk, E. & Hansel, A. (2000). “Cyanobacterial cell walls: News from an unusual prokaryotic envelope”. J. Bacteriol.182: 1191-1199. doi:10.1128/JB.182.5.1191-1199.2000.
^Stewart, I., Schluter, P.J. & Shaw, G.R. (2006). “Cyanobacterial lipopolysaccharides and human health - a review”. Environ Health5: 7. doi:10.1186/1476-069X-5-7.
^Flores, E., Herrero, A., Forchhammer, K. & Maldener, I. (2016). “Septal junctions in filamentous heterocyst-forming Cyanobacteria”. Trends in Microbiology24: 79-82. doi:10.1016/j.tim.2015.11.011.
^Bornikoel, J., Carrión, A., Fan, Q., Flores, E., Forchhammer, K., Mariscal, V., ... & Maldener, I. (2017). “Role of two cell wall amidases in septal junction and nanopore formation in the multicellular cyanobacterium Anabaena sp. PCC 7120”. Frontiers in Cellular and Infection Microbiology7: 386. doi:10.3389/fcimb.2017.00386.
^Mullineaux, C. W., Mariscal, V., Nenninger, A., Khanum, H., Herrero, A., Flores, E. & Adams, D. (2008). “Mechanism of intercellular molecular exchange in heterocyst-forming cyanobacteria”. EMBO J.27: 1299-1308. doi:10.1038/emboj.2008.66.
^Šmarda, J., Šmajs, D., Komrska, J. & Krzyžánek, V. (2002). “S-layers on cell walls of cyanobacteria”. Micron33: 257-277. doi:10.1016/S0968-4328(01)00031-2.
^Ehlers, K. & Oster, G. (2012). “On the mysterious propulsion of Synechococcus”. PLoS One7: e36081. doi:10.1371/journal.pone.0036081.
^Strom, S. L., Brahamsha, B., Fredrickson, K. A., Apple, J. K. & Rodríguez, A. G. (2012). “A giant cell surface protein in Synechococcus WH8102 inhibits feeding by a dinoflagellate predator”. Environmental Microbiology14: 807-816. doi:10.1111/j.1462-2920.2011.02640.x.
^Pereira, S., Zille, A., Micheletti, E., Moradas-Ferreira, P., De Philippis, R. & Tamagnini, P. (2009). “Complexity of cyanobacterial exopolysaccharides: composition, structures, inducing factors and putative genes involved in their biosynthesis and assembly”. FEMS Microbiology Reviews33: 917-941. doi:10.1111/j.1574-6976.2009.00183.x.
^ abPlude, J.L., Parker, D.L., Schommer, O.J., Timmerman, R.J., Hagstrom, S.A., Joers, J.M. & Hnasko, R. (1991). “Chemical characterization of polysaccharide from the slime layer of the cyanobacterium Microcystis flos-aquae C3-40”. Appl. Environ. Microbiol.57: 1696-1700.
^ abDe Philippis, R. & Vincenzini, M. (1998). “Exocellular polysaccharides from cyanobacteria and their possible applications”. FEMS Microbiology Reviews22: 151-175. doi:10.1111/j.1574-6976.1998.tb00365.x.
^De Philippis, R. & Vincenzini, M. (2003). “Outermost polysaccharidic investments of cyanobacteria: nature, significance and possible applications”. Recent Res. Dev. Microbiol.7: 13-22.
^ abMcCarren, J. & Brahamsha, B. (2009). “Swimming motility mutants of marine Synechococcus affected in production and localization of the S-layer protein SwmA”. J. Bacteriol.191: 1111-1114. doi:10.1128/JB.01401-08.
^Reynolds, C. S. (2007). “Variability in the provision and function of mucilage in phytoplankton: facultative responses to the environment”. Hydrobiologia578: 37-45. doi:10.1007/s10750-006-0431-6.
^Ehling-Schulz, M., Bilger, W. & Scherer, S. (1997). “UV-B-induced synthesis of photoprotective pigments and extracellular polysaccharides in the terrestrial cyanobacterium Nostoc commune”. J. Bacteriol.179: 1940-1945. doi:10.1128/jb.179.6.1940-1945.1997.
^Storme, J. Y., Golubic, S., Wilmotte, A., Kleinteich, J., Velázquez, D. & Javaux, E. J. (2015). “Raman characterization of the UV-protective pigment gloeocapsin and its role in the survival of cyanobacteria”. Astrobiology15: 843-857. doi:10.1089/ast.2015.1292.
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