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Prochlorococcus marinus

Prochlorococcus ( 加泰隆語 )

由wikipedia CA提供

Prochlorococcus és un gènere de cianobacteris marins de grandària molt petita (0,6 micres) amb pigmentació inusual (clorofil·la b). Aquests bacteris pertanyen a la picoplàncton fotosintètic i són, probablement, l'organisme fotosintètic més abundant a la Terra. Els microbis del gènere Prochlorococcus estan entre els principals productors primaris de l'oceà, sent els responsables de gran part de la producció fotosintètica d'oxigen.[2][3] L'anàlisi de les seqüències del genoma de 12 soques de Prochlorococcus mostren que 1100 gens són comuns en totes elles, i la grandària mitjana de genoma és d'aproximadament 2.000 gens.[2] En canvi, les algues eucariotes tenen més de 10.000 gens.[4]

Descobriment

Tot i que hi havia diversos registres anteriors d'una petita espècie contenidora de clorofil·la b a l'oceà,[5][6] Prochlorococcus va ser descobert el 1986;[1] l'equip estava format per Sallie W. (Penny) Chisholm de l'Institut de Tecnologia de Massachusetts, Robert J. Olson, de la Institució Oceanogràfica Woods Hole, i altres col·laboradors en el mar dels Sargassos utilitzant la citometria de flux.[7] El primer cultiu de Prochlorococcus es va aïllar en el mar dels Sargassos el 1988 (soca SS120) i al poc temps un altre soca es va obtenir en el mar Mediterrani (soca MED). El nom Prochlorococcus sorgí pel fet que originalment s'havia suposat que Prochlorococcus estava relacionat amb Prochloron i altres bacteris contenidors de clorofil·la b, anomenats procloròfits;[8] però actualment se sap que els procloròfits formen diversos grups filogenètics separats dins del subgrup dels cianobacteris.

L'única espècie del gènere que s'ha descrit és Prochlorococcus marinus.

Morfologia

Les cianobacteris marines són, fins avui, els organismes fotosintètics més petits coneguts; El Prochlorococcus és el més petit presentant només 0,5 a 0,8 micròmetres de diàmetre.[9][3] Aquestes cèl·lules, com totes amb forma de cocos, no tenen mobilitat. Són de vida lliure. Són d'una mida petita, i per tant presenten una alta relació superfície-volum, la qual cosa els dóna un avantatge en aigües pobres en nutrients. Així i tot, se suposa que Prochlorococcus té un requeriment de nutrients molt petit.[10] En general, els individus de Prochlorococcus es divideixen una vegada al dia en la capa subsuperficial o en aigües oligotròfiques.[10]

Distribució

Els Prochlorococcus s'han trobat, de manera abundant, a la zona eufòtica dels oceans tropicals del món.[11] Són, possiblement, les espècies més abundants de la Terra: un sol mil·lilitre d'aigua marina superficial pot contenir 100.000 o més cèl·lules. A nivell mundial, l'abundància mitjana anual és de 2,8 i 3,0 octil·lions (~ 1027 ) d'individus; en comparació, és aproximadament el nombre d'àtoms que conté una tona d'or.[12] Els Prochlorococcus habiten entre el 40° N i 40° S,[10] i dominen a les regions oligotròfiques (pobres en nutrients). Els Prochlorococcus es troben sobretot en un rang de temperatura de 10-33° C i alguns exemplars poden créixer a profunditats amb poca llum (<1 % de llum de la superfície).[2] A aquestes soques se li ha assignat l'ecotips LL (Low Light), a les soques que ocupen menor profunditat en la columna d'aigua se'ls va assignar l'ecotip HL (High light).[13]

Els Prochlorococcus d'ecotip LL presenten una major proporció de clorofil·la b que de clorofil·la a, la qual cosa ajuda a la seva capacitat per absorbir la llum blava.[14] Ja que la llum blava és capaç de penetrar en les aigües de l'oceà més profund que la resta de l'espectre visible, fins a aconseguir profunditats de>200 metres, depenent de la turbidesa de l'aigua. Aquesta profunditat de penetració de la llum blava, combinat amb la capacitat del Prochlorococcus de ecotipo LL d'utilitzar-la per a la fotosíntesi, permet que les poblacions de Prochlorococcus LL sobreviure a profunditats de fins a 200 metres.[15]

Pigments

Els Prochlorococcus està estretament relacionat amb els Synechococcus, un altre cianobacteri fotosintètic abundant, que conté ficobilisomes (antenes per a captació de llum). No obstant, Prochlorochoccus ha evolucionat per utilitzar un únic complex captador de llum, que consisteix predominantment de derivats divinilics de clorofil·la a (Chl a2) i b (Chl b2) i freturosa de clorofil·les monoviniliques i ficobilisomes.[16] Prochlorococcus és l'única espècie silvestre fototròfica oxigènica coneguda que no conté Chl a com a pigment fotosintètic important, i és l'únic procariota conegut amb α-carotè.[17]

Els Prochlorococcus ocupen dos nínxols ecològics diferents, la qual cosa porta a classificar-los en per la seva exposició a la llum, ja sigui baixa (LL) ja sigui alta (HL),[18] que varien en proporcions de pigment (els de l'ecotipus LL té una alta proporció de clorofil·la B2 enfront de la clorofil·la a2 i els de l'ecotipo HL té una baixa relació de b2 enfront de la clorofil·la a2), requeriments de llum, de nitrogen i la utilització de fòsfor, coure, i la sensibilitat per als virus. Aquests "ecotipos" poden diferenciar-se sobre la base de la seqüència dels seus gens d'RNA ribosomal. Les soques adaptats d'alta llum habiten en profunditats entre 25 i 100 m, mentre que les soques adaptats de poca llum habiten en aigües entre 80 i 200 m.[19] Recentment, els genomes de diversos soques de Prochlorococcus s'han pogut seqüenciar.[20][21] Dotze genomes complets s'han seqüenciat que revelen fisiològicament i genèticament diferents llinatges de Prochlorococcus marinus que són 97 % similar en el gen del 16S rRNA.[19]

Vegeu també

Referències

  1. 1,0 1,1 «A novel free-living prochlorophyte occurs at high cell concentrations in the oceanic euphotic zone». Nature, 334, 6180, 1988, pàg. 340–343. DOI: 10.1038/334340a0.
  2. 2,0 2,1 2,2 Munn, C. Marine Microbiology: ecology and applications Second Ed. Garland Science, 2011.
  3. 3,0 3,1 «Life at the Edge of Sight — Scott Chimileski, Roberto Kolter | Harvard University Press» (en en).
  4. «Patterns and Implications of Gene Gain and Loss in the Evolution of Prochlorococcus». PLoS Genetics, 3, 12, desembre 2007, pàg. e231. DOI: 10.1371/journal.pgen.0030231. PMC: 2151091. PMID: 18159947.
  5. «Chroococcoid cyanobacteria in the sea: a ubiquitous and diverse phototrophic biomass». Limnology and Oceanography, 24, 5, 1979, pàg. 928–935. DOI: 10.4319/lo.1979.24.5.0928.
  6. «Unknown chlorophyll a derivatives in the North Sea and the tropical Atlantic Ocean revealed by HPLC analysis». Limnology and Oceanography, 28, 4, 1983, pàg. 757–766. DOI: 10.4319/lo.1983.28.4.0757.
  7. S. W. Chisholm, R. J. Olson, E. R. Zettler, J. Waterbury, R. Goericke & N. Welschmeyer «A novel free-living prochlorophyte occurs at high cell concentrations in the oceanic euphotic zone». Nature, 334, 6180, 1988, pàg. 340–343. DOI: 10.1038/334340a0.
  8. Sallie W. Chisholm, S. L. Frankel, R. Goericke, R. J. Olson, B. Palenik, J. B. Waterbury, L. West-Johnsrud & E. R. Zettler «Prochlorococcus marinus nov. gen. nov. sp.: an oxyphototrophic marine prokaryote containing divinyl chlorophyll a and b». Archives of Microbiology, 157, 3, 1992, pàg. 297–300. DOI: 10.1007/BF00245165.
  9. Biller, Steven J.; Berube, Paul M.; Lindell, Debbie; Chisholm, Sallie W. «Prochlorococcus: the structure and function of collective diversity». Nature Reviews Microbiology, 13, 1, 01-12-2014, pàg. 13–27. DOI: 10.1038/nrmicro3378. PMID: 25435307.
  10. 10,0 10,1 10,2 «Prochlorococcus, a marine photosynthetic prokaryote of global significance.». Microbiology and Molecular Biology Reviews, 63, 1, 1999, pàg. 106–127. PMC: 98958. PMID: 10066832.
  11. Chisholm, S.W.; Frankel, S.; Goericke, R.; Olson, R.; Palenik, B. «Prochlorococcus marinus nov. gen. nov. sp.: an oxyphototrophic marine prokaryote containing divinyl chlorophyll a and b.». Archives of Microbiology, 157, 3, 1992, pàg. 297–300. DOI: 10.1007/bf00245165.
  12. Flombaum, P.; Gallegos, J. L.; Gordillo, R. A.; Rincon, J.; Zabala, L. L. «Present and future global distributions of the marine Cyanobacteria Prochlorococcus and Synechococcus». Proceedings of the National Academy of Sciences, 110, 24, 2013, pàg. 9824–9829. DOI: 10.1073/pnas.1307701110. PMC: 3683724. PMID: 23703908.
  13. Coleman, M.; Sullivan, M.; Martiny, A.; Steglich, C.; Barry, K. «Genomic islands and the ecology and evolution of Prochlorococcus». Science, 311, 5768, 2006, pàg. 1768–1770. DOI: 10.1126/science.1122050. PMID: 16556843.
  14. Ralf, G.; Repeta, D. «The pigments of Prochlorococcus marinus: The presence of divinylchlorophyll a and b in a marine prokaryote». Limnology and Oceanography, 37, 2, 1992, pàg. 425–433. DOI: 10.4319/lo.1992.37.2.0425.
  15. Zinser, E.; Johnson, Z.; Coe, A.; Karaca, E.; Veneziano, D. «Influence of light and temperature on Prochlorococcus ecotype distributions in the Atlantic Ocean». Limnology and Oceanography, 52, 5, 2007, pàg. 2205–2220. DOI: 10.4319/lo.2007.52.5.2205.
  16. «Cyanobacterial photosynthesis in the oceans: the origins and significance of divergent light-harvesting strategies». Trends in Microbiology, 10, 3, 2002, pàg. 134–142. DOI: 10.1016/s0966-842x(02)02319-3.
  17. «The pigments of Prochlorococcus marinus: the presence of divinyl chlorophyll a and b in a marine prokaryote». Limnology and Oceanography, 37, 2, 1992, pàg. 425–433. DOI: 10.4319/lo.1992.37.2.0425.
  18. «Niche-partitioning of Prochlorococcus in a stratified water column in the eastern North Atlantic Ocean». Applied and Environmental Microbiology, 65, 1999, pàg. 2585–2591.
  19. 19,0 19,1 «Taxonomic resolution, ecotypes and biogeography of Prochlorococcus». Environmental Microbiology, 11, 4, 2009, pàg. 823–832. DOI: 10.1111/j.1462-2920.2008.01803.x.
  20. «Genome divergence in two Prochlorococcus ecotypes reflects oceanic niche differentiation» (– Scholar search). Nature, 424, 6952, 2003, pàg. 1042–1047. DOI: 10.1038/nature01947. PMID: 12917642.
  21. «Genome sequence of the cyanobacterium Prochlorococcus marinus SS120, a nearly minimal oxyphototrophic genome». Proceedings of the National Academy of Sciences, 100, 17, 2003, pàg. 10020–10025. DOI: 10.1073/pnas.1733211100. PMC: 187748. PMID: 12917486.

Bibliografia

Enllaços externs

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Prochlorococcus: Brief Summary ( 加泰隆語 )

由wikipedia CA提供

Prochlorococcus és un gènere de cianobacteris marins de grandària molt petita (0,6 micres) amb pigmentació inusual (clorofil·la b). Aquests bacteris pertanyen a la picoplàncton fotosintètic i són, probablement, l'organisme fotosintètic més abundant a la Terra. Els microbis del gènere Prochlorococcus estan entre els principals productors primaris de l'oceà, sent els responsables de gran part de la producció fotosintètica d'oxigen. L'anàlisi de les seqüències del genoma de 12 soques de Prochlorococcus mostren que 1100 gens són comuns en totes elles, i la grandària mitjana de genoma és d'aproximadament 2.000 gens. En canvi, les algues eucariotes tenen més de 10.000 gens.

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Prochlorococcus ( 捷克語 )

由wikipedia CZ提供

Prochlorococcus je rod velmi malých (0.6 µm) mořských sinic, které uvnitř svých buněk obsahují pro sinice neobvyklý chlorofyl b. Řadí se k fotosyntetizujícímu mořskému pikoplanktonu. Je to pravděpodobně nejrozšířenější (počtem odhadovaných jedinců) fotosyntetizující organismus na Zemi vůbec.

Historie výzkumu

Přestože zprávy o existenci sinic s chlorofylem b byly již dříve,[1][2], Prochlorococcus byl popsán až v roce 1986[3] Sallie W. (Penny) Chisholmovou z Massachusettského technologického institutu, Robert J. Olsonem z Woods Hole Oceanographic Institution a jejich dalšími spolupracovníky. Nález pocházel z Sargasového moře a buňky byly prokázány na základě průtokové cytometrie. Jméno Prochlorococcus[4] pochází z domněnky objevitelů, že je tento druh příbuzný již dříve známé sinici rodu Prochloron a dalším sinicím s chlorofylem b (tehdy nazývány prochlorofyty).

Popis

Mořské sinice jsou v současnosti nejmenší známé fotosyntetizující organismy, Prochlorococcus dosahuje velikosti v průměru od 0,5 do 0,8 mikrometrů. Také pravděpodobně představují nejhojněji se vyskytující druh na Zemi,[5][6] jediný mililitr vody jich obsahuje i 100 000 buněk. V celém oceánu jich může být až 100 oktalionů (1029).[7] Nejběžněji se vyskytuje Prochlorococcus mezi 40 stupněm severní šířky a 40 stupněm jižní šířky, a to především v oligotrofních (na živiny chudých) vodách[8] Z celkového množství kyslíku produkovaného fotosyntézou vytvoří tento druh sinic až 20% a rovněž tvoří významnou složku potravního řetězce v oceánu.[9]

V nedávné době byly sekvenovány genomy několika druhů rodu Prochlorococcus.[10][11]

Fotosyntéza

Prochlorococcus má unikátní systém využívání světelného záření, při němž se k absorpci světla používají místo monovinyl chlorofylů takzvané divinyl deriváty chlorofylu. Pokud žije Prochlorococcus ve vodách s dostatkem světla, označují se tyto kmeny jako „high light“ (HL), v opačném případě tedy „low light“ (LL)[12]. Tyto dva ekotypy se liší svými nároky na světlo, poměrem svých pigmentů, využíváním fosforu a dusíku a dokonce náchylností k otravě mědí a k nakažení cyanofágy. Mohou však být rozeznány i na základě sekvencí jejich rRNA genu.

Odkazy

Reference

V tomto článku byl použit překlad textu z článku Prochlorococcus na anglické Wikipedii.

  1. P. W. Johnson & J. M. Sieburth. Chroococcoid cyanobacteria in the sea: a ubiquitous and diverse phototrophic biomass. Limnology and Oceanography. 1979, roč. 24, s. 928–935.
  2. W. W. C. Gieskes & G. W. Kraay. Unknown chlorophyll a derivatives in the North Sea and the tropical Atlantic Ocean revealed by HPLC analysis. Limnology and Oceanography. 1983, roč. 28, s. 757–766.
  3. S. W. Chisholm, R. J. Olson, E. R. Zettler, J. Waterbury, R. Goericke & N. Welschmeyer. A novel free-living prochlorophyte occurs at high cell concentrations in the oceanic euphotic zone. Nature. 1988, roč. 334, s. 340–343. DOI:10.1038/334340a0.
  4. Sallie W. Chisholm, S. L. Frankel, R. Goericke, R. J. Olson, B. Palenik, J. B. Waterbury, L. West-Johnsrud & E. R. Zettler. Prochlorococcus marinus nov. gen. nov. sp.: an oxyphototrophic marine prokaryote containing divinyl chlorophyll a and b. Archives of Microbiology. 1992, roč. 157, s. 297–300. DOI:10.1007/BF00245165.
  5. ROBERT C. KING; WILLIAM D. STANSFIELD; PAMELA K. MULLIGAN. A Dictionary of Genetics, Seventh Edition. [s.l.]: Oxford University Press, 2006.
  6. http://www.scientificamerican.com/article.cfm?id=the-cells-that-rule-the-s
  7. DGF [online]. NASA. Dostupné online.
  8. F. Partensky, W. R. Hess & D. Vaulot. Prochlorococcus, a marine photosynthetic prokaryote of global significance. Microbiology and Molecular Biology Reviews. 1999, roč. 63, s. 106–127.
  9. The Most Important Microbe You've Never Heard Of
  10. G. Rocap, F. W. Larimer, J. Lamerdin, S. Malfatti, P. Chain, N. A. Ahlgren, A. Arellano, M. Coleman, L. Hauser, W. R. Hess, Z. I. Johnson, M. Land, D. Lindell, A. F. Post, W. Regala, M. Shah, S. L. Shaw, C. Steglich, M. B. Sullivan, C. S. Ting, A. Tolonen, E. A. Webb, E. R. Zinser & S. W. Chisholm. Genome divergence in two Prochlorococcus ecotypes reflects oceanic niche differentiation. Nature. 2003, roč. 424, s. 1042–1047. Dostupné v archivu pořízeném dne 2004-12-11. DOI:10.1038/nature01947.
  11. A. Dufresne, M. Salanoubat, F. Partensky, F. Artiguenave, I. M. Axmann, V. Barbe, S. Duprat, M. Y. Galperin, E. V. Koonin, F. Le Gall, K. S. Makarova, M. Ostrowski, S. Oztas, C. Robert, I. B. Rogozin, D. J. Scanlan, N. Tandeau de Marsac, J. Weissenbach, P. Wincker, Y. I. Wolf & W. R. Hess. Genome sequence of the cyanobacterium Prochlorococcus marinus SS120, a nearly minimal oxyphototrophic genome. Proceedings of the National Academy of Sciences. 2003, roč. 100, s. 10020–10025. Dostupné online. DOI:10.1073/pnas.1733211100. PMID 12917486.
  12. N. J. West & D. J. Scanlan. Niche-partitioning of Prochlorococcus in a stratified water column in the eastern North Atlantic Ocean. Applied and Environmental Microbiology. 1999, roč. 65, s. 2585–2591.

Externí odkazy

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Prochlorococcus: Brief Summary ( 捷克語 )

由wikipedia CZ提供

Prochlorococcus je rod velmi malých (0.6 µm) mořských sinic, které uvnitř svých buněk obsahují pro sinice neobvyklý chlorofyl b. Řadí se k fotosyntetizujícímu mořskému pikoplanktonu. Je to pravděpodobně nejrozšířenější (počtem odhadovaných jedinců) fotosyntetizující organismus na Zemi vůbec.

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Prochlorococcus marinus ( 德語 )

由wikipedia DE提供

Prochlorococcus marinus ist die einzige bekannte und wissenschaftlich beschriebene Art der Gattung Prochlorococcus. Es ist ein vorwiegend marin verbreitetes, Photosynthese betreibendes, einzelliges Cyanobakterium. Prochlorococcus gehört zu den kleinsten bekannten photoautotrophen Organismen und damit zum sog. Picoplankton. Aufgrund seiner hohen Konzentration in weiten Bereichen der Ozeane ist er nach aktuellem Forschungsstand das Lebewesen mit der höchsten Individuenanzahl und zugleich das am weitesten verbreitete Lebewesen der Erde und spielt bei der Primärproduktion organischer Stoffe eine besonders große Rolle.[1][2][3]

Merkmale

 src=
Prochlorococcus, kolorierte SEM-Aufnahme

Die Zellen sind mit 0,5 bis 0,8 µm Durchmesser – verglichen mit anderen Cyanobakterien – klein. Sie gehören damit zu den kleinsten bekannten Photosynthese betreibenden Organismen und werden dem Picoplankton zugeordnet.[4] Die Licht absorbierenden Pigmente (Photosynthesepigmente) von Prochlorococcus bestehen hauptsächlich aus Chlorophyll a2 (Chl a2) und b2 (Chl b2), dies sind Divinyl-Derivate der in Pflanzen vorkommenden Chlorophylle a und b. Mono-Vinyl-Chlorophylle kommen jedoch nicht vor.[1]

Das Genom von Prochlorococcus marinus wurde vollständig sequenziert. Die Analyse der Genomsequenzen von 12 Prochlorococcus-Stämmen zeigt, dass 1.100 Gene allen Stämmen gemeinsam sind und die durchschnittliche Genomgröße bei etwa 2.000 Genen liegt.[2] Im Gegensatz dazu haben eukaryotische Algen über 10.000 Gene.[5]

Verbreitung

Prochlorococcus ist zahlenmäßig nach aktuellem Kenntnisstand der häufigste und am weitesten verbreitete Organismus der Erde.[4] Er kommt hauptsächlich in den Ozeanen zwischen den Breitengraden 40° N und 40° S in den oberen 100 bis 150 m vor, und zwar vor allem in nährstoffarmen (oligotrophen) Bereichen mit einer Wassertemperatur von mindestens 10 °C. Dabei erreicht er Konzentrationen von 1·105 bis 3·105 je Milliliter und 1011 bis 1014 je Quadratmeter und stellt einen beträchtlichen Anteil des Bakterioplanktons aller Ozeane dar.[1]

Lebensweise und Ökologie

Als photoautotropher Organismus steht Prochlorococcus am Beginn der Nahrungskette und ist für einen wesentlichen Teil der marinen Primärproduktion verantwortlich. Die Zellen teilen sich unter natürlichen Bedingungen im Mittel einmal täglich. Das bedeutet, dass jeden Tag 50 Prozent der gesamten Prochlorococcus-Biomasse in die marinen Nahrungsnetze eintreten.

Prochlorococcus besiedelt hauptsächlich zwei ökologische Nischen: Neben oberflächennah lebenden Populationen findet man Photosynthese betreibende Zellen auch bis in Tiefen von über 150 Meter. Hier stehen weniger als 1 Prozent der oberflächennahen Lichtintensität zur Verfügung und das Elektromagnetische Spektrum des Lichtes enthält nur noch den Blauanteil. Diese an Schwachlicht adaptierten Zellen verfügen über Antennenpigmente, die auch blaues Licht geringer Intensität absorbieren können und ein Überleben ermöglichen. Entsprechend werden die Bakterien in zwei Gruppen eingeteilt: Mitglieder der low light (LL)-Gruppe besitzen ein höheres Verhältnis von Chlorophyll b2 : a2 als Mitglieder der high light (HL)-Gruppe. Die Gruppen unterscheiden sich außerdem in ihren Stickstoff- und Phosphatanforderungen sowie in ihrer Sensibilität gegenüber Kupferverbindungen und Viren. Hochlichtadaptierte Stämme bewohnen Tiefen zwischen 25 und 100 m, während niedriglichtadaptierte Stämme Gewässer zwischen 80 und 200 m bewohnen. Diese Ökotypen können anhand der Sequenz ihres ribosomalen RNA-Gens unterschieden werden.[6][7]

Forschungsgeschichte

Die frühesten Hinweise auf sehr kleine Chlorophyll-b-haltige Cyanobakterien im Ozean stammen aus den Jahren 1979[8] und 1983.[9] Die Gattung Prochlorococcus wurde 1986 von Sallie W. (Penny) Chisholm (Massachusetts Institute of Technology), Robert J. Olson (Woods Hole Oceanographic Institution) und weiteren Mitarbeitern in der Sargassosee mittels Durchflusszytometrie entdeckt.[10] Für die Entdeckung wurde Chisholm 2019 mit dem Crafoord-Preis ausgezeichnet.[11] Die erste Kultur von Prochlorococcus wurde 1988 in der Sargassosee isoliert (Stamm SS120) und kurz darauf wurde ein weiterer Stamm aus dem Mittelmeer gewonnen (Stamm MED). Der Name Prochlorococcus wurde gewählt, weil man ursprünglich annahm, dass Prochlorococcus mit Prochloron und anderen Chlorophyll b enthaltenden Bakterien, sodass auch die hier verwendete Systematik nach NCBI die Gattung in die Prochlorophyten (Prochlorales) einreihte.[12]

Systematik

 src=
Durchflusszytometrie einer Meerwasserprobe, in der drei Picoplanktongruppen erkennbar sind (Prochlorococcus, Synechococcus und Pico-Eukaryoten)

Inzwischen ist aber bekannt, dass die Prochlorophyten (wiss. Prochlorophyta) mehrere separate phylogenetische Zweige innerhalb der Cyanobakterien bilden. Anhand der rRNA-Sequenzen erkannte man, dass es sich bei Prochlorococcus um eine eigenständige Gruppe unter den Cyanobakterien handelt, die zwar die Divinyl-Derivate von Chlorophyll a und b, aber keine Mono-Vinyl-Chlorophylle enthalten. Prochlorococcus ist der einzige bekannte sauerstoffhaltige phototrophe Wildtyp, der kein Chl a als Hauptphotosynthesepigment enthält, und ist der einzige bekannte Prokaryote mit α-Carotin.[13]

Neuere Phylogenien (Stand 2021) fassen die Gattung Prochlorococcus und die marinen Synechococcus in einer Klade „mariner Picocyanobacteria“ (auch „marine SynPro-Gruppe“ genannt) zusammen, deren letzter gemeinsamer Ahn (LGA oder MRCA) vor etwa 414 (340 bis 419) Millionen Jahren gelebt hat. Die Auseinanderentwicklung (Divergenz) dieser Gruppe und der Gattungen Cyanobium, Aphanothece, sowie anderer verwandter Synechococcus-Vertreter wird im späten Ediacarium (vor 571 Millionen Jahren) angenommen.[14]

Viren

 src=
Adsorption von P-SSP7-Phagen (Prochlorococcus virus PSSP7) an Prochlorococcus marinus MED4, visualisiert durch Kryo-ET.

Es gibt eine Reihe bekannter Viren, die Prochlorococcus infizieren, und daher nicht-taxonomisch als Cyanophagen klassifiziert werden. Die vom International Committee on Taxonomy of Viruses (ICTV) bestätigten Spezies (Stand Januar 2022) gehören den Familien Autographiviridae und Myoviridae (beide in der Klasse Caudoviricetes) an:[15][16]

  • Gattung Banchanvirus
  • Prochlorococcus virus SS120-1
  • Gattung Cheungvirus
  • Prochlorococcus virus NATL1A7
  • Gattung Lingvirus
  • Prochlorococcus virus PGSP1
  • Gattung Tangaroavirus
  • Prochlorococcus virus 951510a
  • Prochlorococcus virus NATL2A133
  • Prochlorococcus virus PSSP10
  • Gattung Tiamatvirus
  • Prochlorococcus virus PSSP7
  • Gattung Tritonvirus
  • Prochlorococcus virus PSSP3
  • Gattung Brizovirus
  • Prochlorococcus virus Syn33
  • Gattung Eurybiavirus
  • Prochlorococcus virus MED4-213
  • Prochlorococcus virus PHM1
  • Prochlorococcus virus PHM2
  • Gattung Libanvirus
  • Prochlorococcus virus PTIM40
  • Gattung Palaemonvirus
  • Prochlorococcus virus PSSM7
  • Gattung Ronodorvirus
  • Prochlorococcus virus PSSM3
  • Prochlorococcus virus PSSM4
  • Gattung Salacisavirus
  • Prochlorococcus virus PSSM2
  • Gattung Vellamovirus
  • Prochlorococcus virus Syn1

Belege

  1. a b c F. Partensky, W. R. Hess, D. Vaulot: Prochlorococcus, a marine photosynthetic prokaryote of global significance, in: Microbiology and Molecular Biology Reviews, Band 63, Nr. 1, 1999, S. 106–127.
  2. a b C. Munn: Marine Microbiology: ecology and applications Second Ed. Garland Science, 2011.
  3. Life at the Edge of Sight — Scott Chimileski, Roberto Kolter | Harvard University Press (en) In: www.hup.harvard.edu.
  4. a b Thomas M. Smith, Robert L. Smith: Ökologie. Pearson Studium, München 2009; S. 447. ISBN 978-3-8273-7313-7.
  5. G. C. Kettler, A. C. Martiny, K. Huang et al.: Patterns and Implications of Gene Gain and Loss in the Evolution of Prochlorococcus. In: PLOS Genetics. 3, Nr. 12, Dezember 2007, S. e231. doi:10.1371/journal.pgen.0030231. PMID 18159947. PMC 2151091 (freier Volltext).
  6. N. J. West, D. J. Scanlan: Niche-partitioning of Prochlorococcus in a stratified water column in the eastern North Atlantic Ocean. In: Applied and Environmental Microbiology. 65, Nr. 6, 1999, S. 2585–2591. doi:10.1128/AEM.65.6.2585-2591.1999. PMID 10347047. PMC 91382 (freier Volltext).
  7. A. C. Martiny, A. Tai, D. Veneziano, F. Primeau, S. Chisholm: Taxonomic resolution, ecotypes and biogeography of Prochlorococcus. In: Environmental Microbiology. 11, Nr. 4, 1. April 2009, S. 823–832. doi:10.1111/j.1462-2920.2008.01803.x. PMID 19021692.
  8. Paul W. Johnson, John McN. Sieburth: Chroococcoid cyanobacteria in the sea: a ubiquitous and diverse phototrophic biomass. In: Limnology and Oceanography. 24, Nr. 5, September 1979, S. 928–935. bibcode:1979LimOc..24..928J. doi:10.4319/lo.1979.24.5.0928.
  9. Winfried W. Gieskes, Gijsbert W. Kraay: Unknown chlorophyll a derivatives in the North Sea and the tropical Atlantic Ocean revealed by HPLC analysis. In: Limnology and Oceanography. 28, Nr. 4, Juli 1983, S. 757–766. bibcode:1983LimOc..28..757G. doi:10.4319/lo.1983.28.4.0757.
  10. Sallie W. Chisholm, Robert J. Olson, Erik R. Zettler, Ralf Goericke, John B. Waterbury, Nicholas A. Welschmeyer: A novel free-living prochlorophyte occurs at high cell concentrations in the oceanic euphotic zone. In: Nature. 334, Nr. 6180, 1. Juli 1988, S. 340–343. bibcode:1988Natur.334..340C. doi:10.1038/334340a0.
  11. The Crafoord Prize in Biosciences 2019, The Royal Swedish Academy of Sciences
  12. Sallie W. Chisholm, S. L. Frankel, R. Goericke, R. J. Olson, B. Palenik, J. B. Waterbury, L. West-Johnsrud & E. R. Zettler: Prochlorococcus marinus nov. gen. nov. sp.: an oxyphototrophic marine prokaryote containing divinyl chlorophyll a and b. In: Archives of Microbiology. 157, Nr. 3, 1992, S. 297–300. doi:10.1007/BF00245165.
  13. R. Goericke, D. Repeta: The pigments of Prochlorococcus marinus: the presence of divinyl chlorophyll a and b in a marine prokaryote. In: Limnology and Oceanography. 37, Nr. 2, 1992, S. 425–433. bibcode:1992LimOc..37..425R. doi:10.4319/lo.1992.37.2.0425.
  14. G. P. Fournier, K. R. Moore, L. T. Rangel, J. G. Payette, L. Momper, T. Bosak: The Archean origin of oxygenic photosynthesis and extant cyanobacterial lineages, Band 288, Nr. 1959, 29. September 2021, doi:10.1098/rspb.2021.0675, PMID 34583585. Siehe insbes. Fig. 2
  15. ICTV: ICTV Master Species List 2020.v1, Email ratification March 2021 (MSL #36)
  16. NCBI: Suche: Prochlorococcus Viren (tendenziell ICTV-bestätigt), und Suche: Prochlorococcus Phagen (tendenziell ICTV-unbestätigt).

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wikipedia DE

Prochlorococcus marinus: Brief Summary ( 德語 )

由wikipedia DE提供

Prochlorococcus marinus ist die einzige bekannte und wissenschaftlich beschriebene Art der Gattung Prochlorococcus. Es ist ein vorwiegend marin verbreitetes, Photosynthese betreibendes, einzelliges Cyanobakterium. Prochlorococcus gehört zu den kleinsten bekannten photoautotrophen Organismen und damit zum sog. Picoplankton. Aufgrund seiner hohen Konzentration in weiten Bereichen der Ozeane ist er nach aktuellem Forschungsstand das Lebewesen mit der höchsten Individuenanzahl und zugleich das am weitesten verbreitete Lebewesen der Erde und spielt bei der Primärproduktion organischer Stoffe eine besonders große Rolle.

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புரோக்குளோரோக்காக்கசு ( 坦米爾語 )

由wikipedia emerging languages提供

புரோக்குளோரோக்காக்கசு என்பது மிகவும் நுண்ணிய, 0.6 மைக்ரோமீட்டர் அளவே உள்ள, வித்தியாசமான கடல்வாழ் கோளவுயிரி ஆகும். இதுவே அநேகமாக உலகில் அதிகமாகக் காணப்படும் ஒளிச்சேர்க்கை உயிரியாக இருக்கலாம்.

புரோக்குளோரோக்காக்கசு முதன்முதலில் 1986ல் கண்டுபிடிக்கப்பட்டது. மாசச்சூசட்சு நுட்பியல் கழகத்தின் சேல்லி (பென்னி) சிசோல்ம் என்பவராலும், வுட்சு ஹோல் ஓசனோகிராபிக் கழகத்தின் ராபர்ட்டு ஜே. ஓல்சன் என்பவராலும், அவர்களுடைய கூட்டுழைப்பாளர்களாலும் சர்காசோக் கடலில் இவ்வுயிரி கண்டுபிடிக்கப்பட்டது. [1]

இவையே இதுவரை அறியப்பட்டதிலேயே மிகவும் சிறிய அளவிலான ஒளிச்சேர்க்கை உயிரிகளாகும். இவற்றின் நீளம் 0.5 முதல் 0.8 மைக்ரோமீட்டர் அளவே இருக்கும். ஒற்றை மில்லி லிட்டர் கடல்நீரில் ஒரு லட்சம் செல்களுக்கும் மேலே இருக்கும். உலகிலேயே மிகவும் அதிகமாகக் காணப்படும் உயிரினம் இதுவாகத் தான் இருக்கும்.

புரோக்குளோரோக்காக்கசின் ஒளிச்சேர்க்கும் பொருள் தனித்தன்மை வாய்ந்தது. 1986க்கும் முன்னர் இவற்றை உலகம் அறிந்திருக்கவில்லை என்றாலும், உலக ஆக்சிஜன் அளவில் ஏறத்தாழ 20 விழுக்காடுக்கு இவையே காரணம் என்று நம்பப்படுகிறது.

உசாத்துணைகள்

  1. S. W. Chisholm, R. J. Olson, E. R. Zettler, J. Waterbury, R. Goericke & N. Welschmeyer (1988). "A novel free-living prochlorophyte occurs at high cell concentrations in the oceanic euphotic zone". Nature 334: 340–343.

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புரோக்குளோரோக்காக்கசு: Brief Summary ( 坦米爾語 )

由wikipedia emerging languages提供

புரோக்குளோரோக்காக்கசு என்பது மிகவும் நுண்ணிய, 0.6 மைக்ரோமீட்டர் அளவே உள்ள, வித்தியாசமான கடல்வாழ் கோளவுயிரி ஆகும். இதுவே அநேகமாக உலகில் அதிகமாகக் காணப்படும் ஒளிச்சேர்க்கை உயிரியாக இருக்கலாம்.

புரோக்குளோரோக்காக்கசு முதன்முதலில் 1986ல் கண்டுபிடிக்கப்பட்டது. மாசச்சூசட்சு நுட்பியல் கழகத்தின் சேல்லி (பென்னி) சிசோல்ம் என்பவராலும், வுட்சு ஹோல் ஓசனோகிராபிக் கழகத்தின் ராபர்ட்டு ஜே. ஓல்சன் என்பவராலும், அவர்களுடைய கூட்டுழைப்பாளர்களாலும் சர்காசோக் கடலில் இவ்வுயிரி கண்டுபிடிக்கப்பட்டது.

இவையே இதுவரை அறியப்பட்டதிலேயே மிகவும் சிறிய அளவிலான ஒளிச்சேர்க்கை உயிரிகளாகும். இவற்றின் நீளம் 0.5 முதல் 0.8 மைக்ரோமீட்டர் அளவே இருக்கும். ஒற்றை மில்லி லிட்டர் கடல்நீரில் ஒரு லட்சம் செல்களுக்கும் மேலே இருக்கும். உலகிலேயே மிகவும் அதிகமாகக் காணப்படும் உயிரினம் இதுவாகத் தான் இருக்கும்.

புரோக்குளோரோக்காக்கசின் ஒளிச்சேர்க்கும் பொருள் தனித்தன்மை வாய்ந்தது. 1986க்கும் முன்னர் இவற்றை உலகம் அறிந்திருக்கவில்லை என்றாலும், உலக ஆக்சிஜன் அளவில் ஏறத்தாழ 20 விழுக்காடுக்கு இவையே காரணம் என்று நம்பப்படுகிறது.

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Prochlorococcus marinus ( 英語 )

由wikipedia EN提供

Prochlorococcus is a genus of very small (0.6 μm) marine cyanobacteria with an unusual pigmentation (chlorophyll a2 and b2). These bacteria belong to the photosynthetic picoplankton and are probably the most abundant photosynthetic organism on Earth. Prochlorococcus microbes are among the major primary producers in the ocean, responsible for a large percentage of the photosynthetic production of oxygen.[1][2] Prochlorococcus strains, called ecotypes, have physiological differences enabling them to exploit different ecological niches.[3] Analysis of the genome sequences of Prochlorococcus strains show that 1,273[4] genes are common to all strains, and the average genome size is about 2,000 genes.[1] In contrast, eukaryotic algae have over 10,000 genes.[4]

Discovery

Although there had been several earlier records of very small chlorophyll-b-containing cyanobacteria in the ocean,[5][6] Prochlorococcus was discovered in 1986[7] by Sallie W. (Penny) Chisholm of the Massachusetts Institute of Technology, Robert J. Olson of the Woods Hole Oceanographic Institution, and other collaborators in the Sargasso Sea using flow cytometry. Chisholm was awarded the Crafoord Prize in 2019 for the discovery.[8] The first culture of Prochlorococcus was isolated in the Sargasso Sea in 1988 (strain SS120) and shortly another strain was obtained from the Mediterranean Sea (strain MED). The name Prochlorococcus[9] originated from the fact it was originally assumed that Prochlorococcus was related to Prochloron and other chlorophyll-b-containing bacteria, called prochlorophytes, but it is now known that prochlorophytes form several separate phylogenetic groups within the cyanobacteria subgroup of the bacteria domain. The only species within the genus described is Prochlorococcus marinus, although two subspecies have been named for low-light and high-light adapted niche variations. [10]

Morphology

Marine cyanobacteria are to date the smallest known photosynthetic organisms; Prochlorococcus is the smallest at just 0.5 to 0.7 micrometres in diameter.[11][2] The coccoid shaped cells are non-motile and free-living. Their small size and large surface-area-to-volume ratio, gives them an advantage in nutrient-poor water. Still, it is assumed that Prochlorococcus have a very small nutrient requirement.[12] Moreover, Prochlorococcus have adapted to use sulfolipids instead of phospholipids in their membranes to survive in phosphate deprived environments.[13] This adaptation allows them to avoid competition with heterotrophs that are dependent on phosphate for survival.[13] Typically, Prochlorococcus divide once a day in the subsurface layer or oligotrophic waters.[12]

Distribution

Prochlorococcus is abundant in the euphotic zone of the world's tropical oceans.[14] It is possibly the most plentiful genus on Earth: a single millilitre of surface seawater may contain 100,000 cells or more. Worldwide, the average yearly abundance is (2.8 to 3.0)×1027 individuals[15] (for comparison, that is approximately the number of atoms in a ton of gold). Prochlorococcus is ubiquitous between 40°N and 40°S and dominates in the oligotrophic (nutrient-poor) regions of the oceans.[12] Prochlorococcus is mostly found in a temperature range of 10-33 °C and some strains can grow at depths with low light (<1% surface light).[1] These strains are known as LL (Low Light) ecotypes, with strains that occupy shallower depths in the water column known as HL (High Light) ecotypes.[16] Furthermore, Prochlorococcus are more plentiful in the presence of heterotrophs that have catalase abilities.[17] Prochlorococcus do not have mechanisms to degrade reactive oxygen species and rely on heterotrophs to protect them.[17] The bacterium accounts for an estimated 13-48% of the global photosynthetic production of oxygen, and forms part of the base of the ocean food chain.[18]

Pigments

Prochlorococcus is closely related to Synechococcus, another abundant photosynthetic cyanobacteria, which contains the light-harvesting antennae phycobilisomes. However, Prochlorochoccus has evolved to use a unique light-harvesting complex, consisting predominantly of divinyl derivatives of chlorophyll a (Chl a2) and chlorophyll b (Chl b2) and lacking monovinyl chlorophylls and phycobilisomes.[19] Prochlorococcus is the only known wild-type oxygenic phototroph that does not contain Chl a as a major photosynthetic pigment, and is the only known prokaryote with α-carotene.[20]

Genome

The genomes of several strains of Prochlorococcus have been sequenced.[21][22] Twelve complete genomes have been sequenced which reveal physiologically and genetically distinct lineages of Prochlorococcus marinus that are 97% similar in the 16S rRNA gene.[23]

The high-light ecotype has the smallest genome (1,657,990 basepairs, 1,716 genes) of any known oxygenic phototroph, but the genome of the low-light type is much larger (2,410,873 base pairs, 2,275 genes).[21]

Ecology

Ancestors of Prochlorococcus contributed to the production of early atmospheric oxygen.[24] Despite Prochlorococcus being one of the smallest types of marine phytoplankton in the world's oceans, its substantial number make it responsible for a major part of the oceans', world's photosynthesis, and oxygen production.[2] The size of Prochlorococcus (0.5 to 0.7 μm)[12] and the adaptations of the various ecotypes allow the organism to grow abundantly in low nutrient waters such as the waters of the tropics and the subtropics (c. 40°N to 40°S);[25] however, they can be found in higher latitudes as high up as 60° north but at fairly minimal concentrations and the bacteria's distribution across the oceans suggest that the colder waters could be fatal. This wide range of latitude along with the bacteria's ability to survive up to depths of 100 to 150 metres, i.e. the average depth of the mixing layer of the surface ocean, allows it to grow to enormous numbers, up to 3×1027 individuals worldwide.[26] This enormous number makes the Prochlorococcus play an important role in the global carbon cycle and oxygen production. Along with Synechococcus (another genus of cyanobacteria that co-occurs with Prochlorococcus) these cyanobacteria are responsible for approximately 50% of marine carbon fixation, making it an important carbon sink via the biological carbon pump (i.e. the transfer of organic carbon from the surface ocean to the deep via several biological, physical and chemical processes).[27] The abundance, distribution and all other characteristics of the Prochlorococcus make it a key organism in oligotrophic waters serving as an important primary producer to the open ocean food webs.

Ecotypes

Prochlorococcus has different "ecotypes" occupying different niches and can vary by pigments, light requirements, nitrogen and phosphorus utilization, copper, and virus sensitivity.[28][11][29] It is thought that Prochlorococcus may occupy potentially 35 different ecotypes and sub-ecotypes within the worlds' oceans. They can be differentiated on the basis of the sequence of the ribosomal RNA gene.[11][28] It has been broken down by NCBI Taxonomy into two different subspecies, Low-light Adapted (LL) or High-light Adapted (HL).[10] There are six clades within each subspecies.[11]

Low-light adapted

Prochlorococcus marinus subsp. marinus is associated with low-light adapted types.[10] It is also further classified by sub-ecotypes LLI-LLVII, where LLII/III has not been yet phylogenetically uncoupled.[11][30] LV species are found in highly iron scarce locations around the equator, and as a result, have lost several ferric proteins.[31] The low-light adapted subspecies is otherwise known to have a higher ratio of chlorophyll b2 to chlorophyll a2,[28] which aids in its ability to absorb blue light.[32] Blue light is able to penetrate ocean waters deeper than the rest of the visible spectrum, and can reach depths of>200 m, depending on the turbidity of the water. Their ability to photosynthesize at a depth where blue light penetrates allows them to inhabit depths between 80 and 200 m.[23][33] Their genomes can range from 1,650,000 to 2,600,000 basepairs in size.[30]

High-light adapted

Prochlorococcus marinus subsp. pastoris is associated with high-light adapted types.[10] It can be further classified by sub-ecotypes HLI-HLVI.[30][11] HLIII, like LV, is also located in an iron-limited environment near the equator, with similar ferric adaptations.[31] The high-light adapted subspecies is otherwise known to have a low ratio of chlorophyll b2 to chlorophyll a2.[28] High-light adapted strains inhabit depths between 25 and 100 m.[23] Their genomes can range from 1,640,000 to 1,800,000 basepairs in size.[30]

Metabolism

Most cyanobacterium are known to have an Incomplete Tricarboxylic Acid Cycle (TCA).[34][35] In this process, 2-oxoglutarate decarboxylase (2OGDC) and succinic semialdehyde dehydrogenase (SSADH), replace the enzyme 2-oxoglutarate dehydrogenase (2-OGDH).[35] Normally, when this enzyme complex joins with NADP+, it can be converted to succinate from 2-oxoglutarate (2-OG).[35] This pathway is non-functional in Prochlorococcus,[35] as succinate dehydrogenase has been lost evolutionarily to conserve energy that may have otherwise been lost to phosphate metabolism.[36]

Strains

Table modified from [30]

See also

References

  1. ^ a b c Munn, C. Marine Microbiology: ecology and applications Second Ed. Garland Science, 2011.
  2. ^ a b c "Life at the Edge of Sight — Scott Chimileski, Roberto Kolter | Harvard University Press". www.hup.harvard.edu. Retrieved 2018-01-26.
  3. ^ Tolonen AC, et al. (October 2006). "Global gene expression of Prochlorococcus ecotypes in response to changes in nitrogen availability". Mol Syst Biol. 2 (53): e53. doi:10.1038/msb4100087. PMC 1682016. PMID 17016519.
  4. ^ a b c d e f g h i j k Kettler GC, Martiny AC, Huang K, et al. (December 2007). "Patterns and Implications of Gene Gain and Loss in the Evolution of Prochlorococcus". PLOS Genetics. 3 (12): e231. doi:10.1371/journal.pgen.0030231. PMC 2151091. PMID 18159947.
  5. ^ P. W. Johnson & J. M. Sieburth (1979). "Chroococcoid cyanobacteria in the sea: a ubiquitous and diverse phototrophic biomass". Limnology and Oceanography. 24 (5): 928–935. Bibcode:1979LimOc..24..928J. doi:10.4319/lo.1979.24.5.0928.
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Prochlorococcus marinus: Brief Summary ( 英語 )

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Prochlorococcus is a genus of very small (0.6 μm) marine cyanobacteria with an unusual pigmentation (chlorophyll a2 and b2). These bacteria belong to the photosynthetic picoplankton and are probably the most abundant photosynthetic organism on Earth. Prochlorococcus microbes are among the major primary producers in the ocean, responsible for a large percentage of the photosynthetic production of oxygen. Prochlorococcus strains, called ecotypes, have physiological differences enabling them to exploit different ecological niches. Analysis of the genome sequences of Prochlorococcus strains show that 1,273 genes are common to all strains, and the average genome size is about 2,000 genes. In contrast, eukaryotic algae have over 10,000 genes.

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Prochlorococcus ( 瑞典語 )

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Prochlorococcus är en väldigt liten havslevande cyanobakterie med en ovanlig pigmentering, klorofyll b. Den är den minsta och till antalet rikligast förekommande fotosyntetiserande organismen på jorden.

Referenser

Den här artikeln är helt eller delvis baserad på material från engelskspråkiga Wikipedia, Prochlorococcus, 9 januari 2013.
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Prochlorococcus: Brief Summary ( 瑞典語 )

由wikipedia SV提供

Prochlorococcus är en väldigt liten havslevande cyanobakterie med en ovanlig pigmentering, klorofyll b. Den är den minsta och till antalet rikligast förekommande fotosyntetiserande organismen på jorden.

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