Cyanobacterial morphology - Knowledge (XXG)

708: 764: 292: 1234: 853: 423:. The precise molecular circuits that govern those morphological changes are yet to be identified, however, a so-far constant factor is that the cell shape is determined by the rigid PG sacculus which consists of glycan strands crosslinked by peptides. To grow, cells must synthesize new PG while breaking down the existent polymer to insert the newly synthesized material. How cells grow and elongate has been extensively reviewed in model organisms of both, rod-shaped and coccoid bacteria. The molecular basis for morphological plasticity and pleomorphism in more complex bacteria, however, is slowly being elucidated as well. 667: 1298: 1000: 1088: 951: 915: 508: 585: 31: 968: 553: 723: 690: 740: 1252: 899: 939: 524: 1202:, Camargue, France migrate to the upper layer of the mat during the day and are spread homogenously through the mat at night. An in vitro experiment using P. uncinatum also demonstrated this species' tendency to migrate in order to avoid damaging radiation. These migrations are usually the result of some sort of photomovement, although other forms of taxis can also play a role. 4160: 3813: 1551: 1426: 1137:; however, many filamentous species move on surfaces by gliding, a form of locomotion where no physical appendages are seen to aid movement. The actual mechanism behind gliding is not fully understood, although over a century has elapsed since its discovery. One theory suggests that gliding motion in cyanobacteria is mediated by the continuous secretion of 494:, that travel away from the main biomass to bud and form new colonies elsewhere. The cells in a hormogonium are often thinner than in the vegetative state, and the cells on either end of the motile chain may be tapered. To break away from the parent colony, a hormogonium often must tear apart a weaker cell in a filament, called a necridium. 1337:. After a few hours, the trichomes move away from the darker areas onto the lighter areas, forming a photographic positive on the culture. The experiment demonstrates that photomovement is effective not just for discrete light traps, but for minutely patterned, continuously differentiated light fields as well. 1198:

sp. and Spirulina subsalsa found in the hypersaline benthic mats of Guerrero Negro, Mexico migrate downwards into the lower layers during the day in order to escape the intense sunlight and then rise to the surface at dusk. In contrast, the population of Microcoleus chthonoplastes found in hypersaline mats at

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The cells appear to coordinate their gliding direction by an electrical potential that establishes polarity in the trichomes, and thus establishes a "head" and the "tail". Trichomes usually reverse their polarity randomly with an average period on the order of minutes to hours. Many species also form

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Cyanobacteria are ubiquitous, finding habitats in most water bodies and in extreme environments such as the polar regions, deserts, brine lakes and hot springs. They have also evolved surprisingly complex collective behaviours that lie at the boundary between single-celled and multicellular life. For

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are modeled as thin flexible rods that are discretized into sequences of 50 μm edges. Each edge is loaded with a linear spring. (B) The local bending moment is a function of the radius of curvature. (C) Trichomes can glide along their long axis and reverse their direction of movement photophobically.

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In aquatic habitats, unicellular cyanobacteria are considered as an important group regarding abundance, diversity, and ecological character. Unicellular cyanobacteria have spherical, ovoid, or cylindrical cells that may aggregate into irregular or regular colonies bound together by the mucous matrix

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Filamentous cyanobacteria that live in microbial mats often migrate vertically and horizontally within the mat in order to find an optimal niche that balances their light requirements for photosynthesis against their sensitivity to photodamage. For example, the filamentous cyanobacteria Oscillatoria

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is the biological process that causes an organism to develop its shape. Cyanobacteria show a high degree of morphological diversity and can undergo a variety of cellular differentiation processes in order to adapt to certain environmental conditions. This helps them thrive in almost every habitat on

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to produce a wide range of chemicals, including biofuels like biodiesel and ethanol. However, despite their importance to the history of life on Earth, and their commercial and environmental potentials, there remain basic questions of how filamentous cyanobacteria move, respond to their environment

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patterns. Similar patterns have been observed in fossil records. For filamentous cyanobacteria, the mechanics of the filaments is known to contribute to self-organization, for example in determining how one filament will bend when in contact with other filaments or obstacles. Further, biofilms and

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morphologies are tasks that would require active cell wall remodelling and thus far no genes attributed to the different morphotypes have been identified in cyanobacteria. Therefore, the most likely scenario is that genes or their products are differentially regulated during these cell morphology

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Shih, Patrick M.; Wu, Dongying; Latifi, Amel; Axen, Seth D.; Fewer, David P.; Talla, Emmanuel; Calteau, Alexandra; Cai, Fei; Tandeau de Marsac, Nicole; Rippka, Rosmarie; Herdman, Michael; Sivonen, Kaarina; Coursin, Therese; Laurent, Thierry; Goodwin, Lynne; Nolan, Matt; Davenport, Karen W.; Han,

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One factor which can drive morphological changes in cyanobacteria is light. As cyanobacteria are bacteria that use light to fuel their energy-producing photosynthetic machinery they depend on perceiving light in order to optimize their response and to avoid harmful light that could result in the

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Zhu, Zaichun; Piao, Shilong; Myneni, Ranga B.; Huang, Mengtian; Zeng, Zhenzhong; Canadell, Josep G.; Ciais, Philippe; Sitch, Stephen; Friedlingstein, Pierre; Arneth, Almut; Cao, Chunxiang; Cheng, Lei; Kato, Etsushi; Koven, Charles; Li, Yue; Lian, Xu; Liu, Yongwen; Liu, Ronggao; Mao, Jiafu; Pan,

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Dagan, Tal; Roettger, Mayo; Stucken, Karina; Landan, Giddy; Koch, Robin; Major, Peter; Gould, Sven B.; Goremykin, Vadim V.; Rippka, Rosmarie; Tandeau de Marsac, Nicole; Gugger, Muriel; Lockhart, Peter J.; Allen, John F.; Brune, Iris; Maus, Irena; Pühler, Alfred; Martin, William F. (2012-12-07).

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Cyanobacteria have strict light requirements. Too little light can result in insufficient energy production, and in some species may cause the cells to resort to heterotrophic respiration. Too much light can inhibit the cells, decrease photosynthesis efficiency and cause damage by bleaching. UV

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Cellular functions require a well-organized and coordinated internal structure to operate effectively. Cells need to build, sustain, and sometimes modify their shape, which allows them to rapidly change their behaviour in response to external factors. During different life cycle stages, such as

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and extrudes slime at an acute angle. The sets extrude slime in opposite directions and so only one set is likely to be activated during gliding. An alternative hypothesis is that the cells use contractive elements that produce undulations running over the surface inside the slime tube like an

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trichomes. Considering this complex morphology, it was postulated that certain subsection V-specific (cytoskeletal) proteins could be responsible for this phenotype. However, no specific gene was identified whose distribution was specifically correlated with the cell morphology among different

231:(PG) layer between the inner and outer membrane, thus containing features of both Gram phenotypes. Additionally, the degree of PG crosslinking is much higher in cyanobacteria than in other Gram-negative bacteria, although teichoic acids, typically present in Gram-positive bacteria, are absent. 105:

by which a trichome modulates its gliding according to the incident light. The latter has been found to play an important role in guiding the trichomes to optimal lighting conditions, which can either inhibit the cells if the incident light is too weak, or damage the cells if too strong.

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transitions, as it has been hypothesized for most bacteria. In multicellular cyanobacteria, division of labor between cells within a trichome is achieved by different cell programing strategies. Thus, gene regulation occurs differentially in these specific cell types .

238:, their cellular morphologies are extremely diverse and range from unicellular species to complex cell-differentiating, multicellular species. Based on this observation, cyanobacteria have been classically divided into five subsections. Subsection I cyanobacteria ( 1172:

Through collective interaction, filamentous cyanobacteria self-organize into colonies or biofilms, symbiotic communities found in a wide variety of ecological niches. Their larger-scale collective structures are characterized by diverse shapes including bundles,

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Palinska, Katarzyna A.; Liesack, Werner; Rhiel, Erhard; Krumbein, W. E. (17 October 1996). "Phenotype variability of identical genotypes: the need for a combined approach in cyanobacterial taxonomy demonstrated on Merismopedia-like isolates".

405:, or the ability of one cell to alternate between different shapes, is a common strategy of many bacteria in response to environmental changes or as part of their normal life cycle. Bacteria may alter their shape by simpler transitions from 763: 283:

cyanobacterial subsections. Therefore, it seems more likely that differential expression of cell growth and division genes rather than the presence or absence of a single gene is responsible for the cyanobacterial morphological diversity.

707: 967: 212:. Constant influx of new findings finally established that numerous prokaryotic cellular functions, including cell division, cell elongation or bacterial microcompartment segregation are governed by the prokaryotic cytoskeleton. 40:(A) spherical and ovoid unicellular, (B) colonial, (C) filamentous, (D) spiral, (E) unsheathed trichome, (F) sheathed trichome, (G) false branching, (H) true branching, (I) different cell types in filamentous cyanobacteria. 3162:

Yaozhong; Peng, Shushi; Peñuelas, Josep; Poulter, Benjamin; Pugh, Thomas A. M.; Stocker, Benjamin D.; Viovy, Nicolas; Wang, Xuhui; Wang, Yingping; Xiao, Zhiqiang; Yang, Hui; Zaehle, Sönke; Zeng, Ning (2016-04-25).

1016: 198:, a bacterial actin homolog. These discoveries started an intense search for other cytoskeletal proteins in bacteria and archaea which finally led to the identification of bacterial IF-like proteins such as 1213:. Gliding in filamentous cyanobacteria appears to be powered by a "slime jet" mechanism, in which the cells extrude a gel that expands quickly as it hydrates providing a propulsion force, although some 3667:

Nürnberg, Dennis J.; Mariscal, Vicente; Parker, Jamie; Mastroianni, Giulia; Flores, Enrique; Mullineaux, Conrad W. (2014). "Branching and intercellular communication in the Section V cyanobacterium

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Walter, M.R.; Bauld, J.; Brock, T.D. (1976). "Chapter 6.2 Microbiology and Morphogenesis of Columnar Stromatolites (Conophyton, Vacerrilla) from Hot Springs in Yellowstone National Park".

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is a form of cell movement that differs from crawling or swimming in that it does not rely on any obvious external organ or change in cell shape and it occurs only in the presence of a

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Farrokh, Parisa; Sheikhpour, Mojgan; Kasaeian, Alibakhsh; Asadi, Hassan; Bavandi, Roya (2019). "Cyanobacteria as an eco-friendly resource for biofuel production: A critical review".

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Meeks JC, Elhai J, Thiel T, Potts M, Larimer F, Lamerdin J, Predki P, Atlas R (2001). "An overview of the genome of Nostoc punctiforme, a multicellular, symbiotic cyanobacterium".

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appear as long thin curved filaments. (b) When rendered inactive, for example by being briefly cooled, the same filaments adopt a more random shape. (c) Under higher magnification

183:(IFs), although other cytoskeletal classes have been identified in recent years. Only the collaborative work of all three cytoskeletal systems enables proper cell mechanics. 1267:

a semi-rigid sheath that is left behind as a hollow tube as the trichome moves forward. When the trichome reverses direction, it can move back into the sheath or break out.

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that can take a multitude of forms. Of particular interest among the many species of cyanobacteria are those that live colonially in elongate hair-like structures, known as

419:, by more complex transitions while establishing multicellularity or by the development of specialized cells, structures or appendages where the population presents a 5495:

Sumner, Dawn Y. (1997). "Late Archean Calcite-Microbe Interactions: Two Morphologically Distinct Microbial Communities That Affected Calcite Nucleation Differently".

1125:. These large colonies provide a rigid, stable and long-term environment for their communities of bacteria. In addition, cyanobacteria-based biofilms can be used as 1102:(D) Trichome collisions are defined between edge-vertex pairs. A vertex that penetrates an edge's volume is repulsed by equal and opposite forces between the pair. 5444:

Allwood, Abigail C.; Walter, Malcolm R.; Kamber, Balz S.; Marshall, Craig P.; Burch, Ian W. (2006). "Stromatolite reef from the Early Archaean era of Australia".

3881:"A Putative O-Linked β-N-Acetylglucosamine Transferase Is Essential for Hormogonium Development and Motility in the Filamentous Cyanobacterium Nostoc punctiforme" 1690: 4529:

Wharton, Robert A.; Parker, Bruce C.; Simmons, George M. (1983). "Distribution, species composition and morphology of algal mats in Antarctic dry valley lakes".

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The long-lasting dogma that prokaryotes, based on their simple cell shapes, do not require cytoskeletal elements was finally abolished by the discovery of

1015: 914: 487:(reproductive, motile filaments). These, together with the intercellular connections they possess, are considered the first signs of multicellularity. 3291:"Growth phase-regulated expression of bolA and morphology of stationary-phase Escherichia coli cells are controlled by the novel sigma factor sigma S" 3832:"Genetic characterization of the hmp locus, a chemotaxis-like gene cluster that regulates hormogonium development and motility in Nostoc punctiforme" 950: 5387:"Resolving MISS conceptions and misconceptions: A geological approach to sedimentary surface textures generated by microbial and abiotic processes" 3931:

Dvořák, Petr; Casamatta, Dale A.; Hašler, Petr; Jahodářová, Eva; Norwich, Alyson R.; Poulíčková, Aloisie (2017). "Diversity of the Cyanobacteria".

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radiation is especially deadly for cyanobacteria, with normal solar levels being significantly detrimental for these microorganisms in some cases.

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Kühn, Juliane; Briegel, Ariane; Mörschel, Erhard; Kahnt, Jörg; Leser, Katja; Wick, Stephanie; Jensen, Grant J; Thanbichler, Martin (2009-12-03).

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Wiltbank, Lisa B.; Kehoe, David M. (2018-11-08). "Diverse light responses of cyanobacteria mediated by phytochrome superfamily photoreceptors".

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earthworm. The trichomes rotate in a spiral fashion, the angle of which corresponds with the pitch angle of Castenholz's contractile trichomes.

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Jones, B.; Renaut, R. W.; Rosen, M. R.; Ansdell, K. M. (2002). "Coniform Stromatolites from Geothermal Systems, North Island, New Zealand".

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Khayatan B, Bains DK, Cheng MH, Cho YW, Huynh J, Kim R, Omoruyi OH, Pantoja AP, Park JS, Peng JK, Splitt SD, Tian MY, Risser DD (May 2017).

722: 645:) secreted during the growth of the colony. Based on the species, the number of cells in each colony may vary from two to several thousand. 335:, it does not necessarily mean it is essential in all other cyanobacteria. N/A indicates that no mutant phenotypes have been described. WT: 291: 1233: 1157:

would lead to motion, with some suggesting they retract, while others suggest they push, to generate forces. Other scholars have suggested

852: 2212:"Bactofilins, a ubiquitous class of cytoskeletal proteins mediating polar localization of a cell wall synthase in Caulobacter crescentus" 376: 1277:

in its movement. Filaments in colonies slide back and forth against each other until the whole mass is reoriented to its light source.

5074:"Evidence that a modified type IV pilus-like system powers gliding motility and polysaccharide secretion in filamentous cyanobacteria" 4008: 785: 402: 5985:"The junctional pore complex, a prokaryotic secretion organelle, is the molecular motor underlying gliding motility in cyanobacteria" 4869:"The junctional pore complex, a prokaryotic secretion organelle, is the molecular motor underlying gliding motility in cyanobacteria" 3724:

Herrero, Antonia; Stavans, Joel; Flores, Enrique (2016). "The multicellular nature of filamentous heterocyst-forming cyanobacteria".

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Videau, Patrick; Rivers, Orion S.; Ushijima, Blake; Oshiro, Reid T.; Kim, Min Joo; Philmus, Benjamin; Cozy, Loralyn M. (2016-04-15).

6371: 4462: 3973: 3651: 3619: 2930: 1584: 999: 2728:"Genomes of Stigonematalean Cyanobacteria (Subsection V) and the Evolution of Oxygenic Photosynthesis from Prokaryotes to Plastids" 739: 278:) that are surrounded by a common sheath, subsection V can produce lateral branches and/or divide in multiple planes, establishing 5809:"Vertical migration of phototrophic bacterial populations in a hypersaline microbial mat from Salins-de-Giraud (Camargue, France)" 5807:

Fourã§Ans, Aude; Solã©, Antoni; Diestra, Ella; Ranchou-Peyruse, Anthony; Esteve, Isabel; Caumette, Pierre; Duran, Robert (2006).

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Egan, Alexander J. F.; Errington, Jeff; Vollmer, Waldemar (2020-05-18). "Regulation of peptidoglycan synthesis and remodelling".

6093:"The role of an alternative sigma factor in motility and pilus formation in the cyanobacterium Synechocystis sp. Strain PCC6803" 5018:"Genetic characterization of thehmplocus, a chemotaxis-like gene cluster that regulates hormogonium development and motility in 3013:

Larkum, A. W. D.; Ritchie, R. J.; Raven, J. A. (2018). "Living off the Sun: Chlorophylls, bacteriochlorophylls and rhodopsins".

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de Boer, Piet; Crossley, Robin; Rothfield, Lawrence (1992). "The essential bacterial cell-division protein FtsZ is a GTPase".

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are capable of a waving motion; the filament oscillates back and forth. In water columns, some cyanobacteria float by forming

681:. Species in this genus divide in only two directions, creating a characteristic grid-like pattern arranged in rows and flats. 262:) are multicellular, cell differentiating cyanobacteria that form specialized cell types in the absence of combined nitrogen ( 781: 391:

growth up to a wavelength of 750 nm. To sense the light across this range of wavelengths, cyanobacteria possess various

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Alberts, Bruce; Heald, Rebecca; Johnson, Alexander; Morgan, David Owen; Raff, Martin C.; Roberts, K.; Walter, Peter (2022).

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Aguilera, Anabella; Klemenčič, Marina; Sueldo, Daniela J.; Rzymski, Piotr; Giannuzzi, Leda; Martin, María Victoria (2021).

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Wolk, C. Peter; Ernst, Annaliese; Elhai, Jeff (1994). "Heterocyst Metabolism and Development". In Donald A. Bryant (ed.).

5123:"PilB localization correlates with the direction of twitching motility in the cyanobacterium Synechocystis sp. PCC 6803" 1282: 1107: 438:

among cyanobacterial taxa, which can also vary within a given strain during its life cycle. Changes in cellular or even

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Halfen, Lawrence N.; Castenholz, Richard W. (1971). "Gliding Motility in the Blue-Green Alga Oscillatoria Princeps 1".

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Halfen, Lawrence N.; Castenholz, Richard W. (1971). "Gliding Motility in the Blue-Green Alga Oscillatoria Princeps 1".

6242:"Enhanced Model for Photophobic Responses of the Blue-Green Alga, <italic>Phormidium uncinatum</italic>". 1117:

example, filamentous cyanobacteria live in long chains of cells that bundle together into larger structures including

938: 420: 5858:"Diel Vertical Movements of the Cyanobacterium Oscillatoria terebriformis in a Sulfide-Rich Hot Spring Microbial Mat" 5649:"Modeling Filamentous Cyanobacteria Reveals the Advantages of Long and Fast Trichomes for Optimizing Light Exposure" 1501:"Modeling Filamentous Cyanobacteria Reveals the Advantages of Long and Fast Trichomes for Optimizing Light Exposure" 4024: 1331:. In Häder's cyanograph experiment a photographic negative is projected onto a Petri dish containing a culture of 2029:"An ATPase domain common to prokaryotic cell cycle proteins, sugar kinases, actin, and hsp70 heat shock proteins" 1778:

Springstein, Benjamin L.; Nürnberg, Dennis J.; Weiss, Gregor L.; Pilhofer, Martin; Stucken, Karina (2020-12-17).

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Wagstaff, James; Löwe, Jan (2018-01-22). "Prokaryotic cytoskeletons: protein filaments organizing small cells".

246:) are also unicellular but can undergo multiple fission events, giving rise to many small daughter cells termed 78:. These filamentous species can contain hundreds to thousands of cells. They often dominate the upper layers of 4924:

Craig, Lisa; Pique, Michael E.; Tainer, John A. (2004). "Type IV pilus structure and bacterial pathogenicity".

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Cliff S.; Rubin, Edward M.; Eisen, Jonathan A.; Woyke, Tanja; Gugger, Muriel; Kerfeld, Cheryl A. (2012-12-31).

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cover the lighter areas of the projection while uncovering the darker areas producing a photographic positive.

1169:. Recent work also suggests that shape fluctuations and capillary forces could be involved in gliding motion. 3104:

Claessen, Dennis; Rozen, Daniel E.; Kuipers, Oscar P.; Søgaard-Andersen, Lotte; Van Wezel, Gilles P. (2014).

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McBride, Mark J. (2001). "Bacterial Gliding Motility: Multiple Mechanisms for Cell Movement over Surfaces".

3771:"Cell Death in Cyanobacteria: Current Understanding and Recommendations for a Consensus on Its Nomenclature" 2846:

Koch, Robin; Kupczok, Anne; Stucken, Karina; Ilhan, Judith; Hammerschmidt, Katrin; Dagan, Tal (2017-08-31).

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Rippka, Rosmarie; Stanier, Roger Y.; Deruelles, Josette; Herdman, Michael; Waterbury, John B. (1979-03-01).

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Toxic cyanobacteria in water : a guide to their public health consequences, monitoring, and management

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Halfen, Lawrence N.; Castenholz, Richard W. (1970). "Gliding in a Blue–Green Alga: A Possible Mechanism".

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Häder, Donat-P. (1987). "EFFECTS OF UV-B IRRADIATION ON PHOTOMOVEMENT IN THE DESMID, Cosmarium cucumis".

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van den Ent, Fusinita; Amos, Linda A.; Löwe, Jan (2001). "Prokaryotic origin of the actin cytoskeleton".

898: 2371:"Mutation of the murC and murB Genes Impairs Heterocyst Differentiation in Anabaena sp. Strain PCC 7120" 392: 379:), but some cyanobacteria may expand on PAR by not only absorbing in the visible spectrum, but also the 180: 5121:

Schuergers, Nils; Nürnberg, Dennis J.; Wallner, Thomas; Mullineaux, Conrad W.; Wilde, Annegret (2015).

2956:"Production of Reactive Oxygen Species by Photosystem II as a Response to Light and Temperature Stress" 1850:

Bi, Erfei; Lutkenhaus, Joe (1991). "FtsZ ring structure associated with division in Escherichia coli".

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Rippka, Rosmarie; Stanier, Roger Y.; Deruelles, Josette; Herdman, Michael; Waterbury, John B. (1979).

3130: 2848:"Plasticity first: molecular signatures of a complex morphological trait in filamentous cyanobacteria" 802:

vegetative cells – the normal, photosynthetic cells that are formed under favorable growing conditions

30: 6395: 6208: 6104: 6049: 5996: 5869: 5820: 5763: 5722: 5660: 5610: 5551: 5504: 5453: 5421: 5398: 5338: 5279: 5222: 5171: 4880: 4723: 4538: 4487: 4059: 3175: 2859: 2620: 2496: 2103: 2040: 1977: 1918: 1859: 1791: 1512: 930: 130: 48: 4969:"Molecular Analysis of Genes in Nostoc punctiforme Involved in Pilus Biogenesis and Plant Infection" 3512: 364: 2267:

Lin, Lin; Thanbichler, Martin (2013). "Nucleotide-independent cytoskeletal scaffolds in bacteria".

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Löwe, Jan; Amos, Linda A. (1998). "Crystal structure of the bacterial cell-division protein FtsZ".

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Hoiczyk, E. (2000). "Gliding motility in cyanobacteria: Observations and possible explanations".

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Hoiczyk, E. (2000). "Gliding motility in cyanobacteria: Observations and possible explanations".

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to multicellular filamentous forms. Their cell size varies from less than 1 μm in diameter (

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is mainly blue-green or brown-green and is commonly found in watering-troughs. It reproduces by

552: 242:) are unicellular and divide by binary fission or budding, whereas subsection II cyanobacteria ( 90:, deserts and polar regions, as well as being widely distributed in more mundane environments. 6419: 6411: 6367: 6181: 6132: 6065: 6014: 5965: 5930: 5895: 5838: 5789: 5688: 5626: 5579: 5469: 5356: 5307: 5256:

Tchoufag, Joël; Ghosh, Pushpita; Pogue, Connor B.; Nan, Beiyan; Mandadapu, Kranthi K. (2019).

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Schulz-Vogt, Heide N; Angert, Esther R; Garcia-Pichel, Ferran (2007-09-28), "Giant Bacteria",

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Gaysina, Lira A.; Saraf, Aniket; Singh, Prashant (2019). "Cyanobacteria in Diverse Habitats".

2895: 2877: 2825: 2807: 2786:"Improving the coverage of the cyanobacterial phylum using diversity-driven genome sequencing" 2765: 2747: 2702: 2656: 2638: 2589: 2571: 2532: 2514: 2465: 2447: 2408: 2390: 2343: 2335: 2292: 2284: 2249: 2231: 2184: 2176: 2127: 2119: 2076: 2058: 2001: 1993: 1942: 1934: 1883: 1875: 1827: 1809: 1733: 1725: 1705: 1672: 1662: 1639: 1621: 1580: 1540: 1456: 1415: 1397: 1041: 820: 747: 697: 657: 560: 460: 3998: 1572: 6403: 6359: 6317: 6274: 6247: 6216: 6171: 6163: 6122: 6112: 6057: 6004: 5957: 5922: 5885: 5877: 5828: 5779: 5771: 5730: 5678: 5668: 5618: 5569: 5559: 5512: 5461: 5416: 5406: 5346: 5297: 5287: 5230: 5179: 5134: 5085: 5033: 4988: 4980: 4933: 4888: 4841: 4812: 4731: 4694: 4647: 4610: 4569: 4546: 4495: 4450: 4417: 4369: 4328: 4320: 4279: 4271: 4229: 4186: 4139: 4123: 4067: 3936: 3900: 3892: 3843: 3792: 3782: 3741: 3733: 3688: 3680: 3639: 3607: 3574: 3527: 3469: 3432: 3416: 3375: 3359: 3318: 3302: 3261: 3245: 3191: 3183: 3125: 3117: 3106:"Bacterial solutions to multicellularity: A tale of biofilms, filaments and fruiting bodies" 3057: 3022: 2985: 2967: 2918: 2885: 2867: 2815: 2797: 2755: 2739: 2692: 2646: 2628: 2579: 2563: 2522: 2504: 2455: 2439: 2398: 2382: 2327: 2276: 2239: 2223: 2166: 2111: 2066: 2048: 1985: 1926: 1867: 1817: 1799: 1717: 1629: 1613: 1530: 1520: 1448: 1405: 1387: 1206: 1142: 905: 456: 415: 406: 98: 71: 331:

in the respective organism. While one gene can be essential in one cyanobacterial organism/

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biomats show some remarkably conserved macro-mechanical properties, typically behaving as

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Drews G. (1959) "Beitröge zur Kenntnis der phototaktischen Reaktionen der Cyanophyceen".

4398: 227:. However, unlike other Gram-negative bacteria, cyanobacteria contain an unusually thick 17: 6399: 6386:

Hangarter, Roger P.; Gest, Howard (2004). "Pictorial Demonstrations of Photosynthesis".

6212: 6108: 6053: 6000: 5961: 5873: 5824: 5767: 5726: 5664: 5614: 5598: 5555: 5508: 5457: 5402: 5342: 5283: 5226: 5175: 4884: 4727: 4683:"Generic Assignments, Strain Histories and Properties of Pure Cultures of Cyanobacteria" 4542: 4491: 4063: 3179: 2863: 2681:"Generic Assignments, Strain Histories and Properties of Pure Cultures of Cyanobacteria" 2624: 2500: 2107: 2044: 1981: 1922: 1863: 1795: 1516: 1225:

have two sets of pores for extruding slime. Each set is organized in a ring at the cell

6322: 6301: 6278: 6251: 6220: 5735: 5710: 5683: 5648: 5574: 5539: 5302: 5257: 5234: 4993: 4968: 4144: 4111: 3905: 3880: 3797: 3770: 3437: 3404: 3380: 3347: 3266: 3233: 2990: 2955: 2922: 2890: 2847: 2820: 2785: 2760: 2727: 2651: 2608: 2527: 2484: 2403: 2370: 2244: 2211: 1822: 1779: 1634: 1601: 1535: 1500: 1410: 1375: 1251: 1138: 958: 926: 795: 372: 328: 259: 243: 161: 6176: 6151: 6009: 5984: 5890: 5857: 5784: 5751: 4893: 4868: 4454: 4333: 4308: 4284: 4259: 4233: 3323: 3290: 2460: 2427: 2171: 2154: 359:

and subsequently in their death. Optimal light conditions may be defined by quantity (

6445: 6127: 6092: 5833: 5808: 5351: 5326: 4667: 4515: 3555: 3513:"How to get (a)round: mechanisms controlling growth and division of coccoid bacteria" 3497: 3403:

Typas, Athanasios; Banzhaf, Manuel; Gross, Carol A.; Vollmer, Waldemar (2011-12-28).

3306: 3085: 2940: 2584: 2551: 2071: 2028: 1153:, as the driving engines of motion. However, it is not clear how the action of these 983: 791: 380: 343: 317: 311: 296: 239: 228: 142: 126: 79: 52: 6286: 6228: 6167: 6026: 5881: 5775: 5430: 5411: 5386: 5368: 5242: 5199: 5055: 4953: 4910: 4853: 4591: 3865: 3710: 3147: 2567: 2443: 2304: 2196: 1745: 648:

Each individual cell (each single cyanobacterium) typically has a thick, gelatinous

6431: 6363: 6077: 5481: 5385:

Davies, Neil S.; Liu, Alexander G.; Gibling, Martin R.; Miller, Randall F. (2016).

5107: 4751: 4275: 4206: 4087: 3611: 3569:

Whitton, Brian A. (1992). "Diversity, Ecology, and Taxonomy of the Cyanobacteria".

3405:"From the regulation of peptidoglycan synthesis to bacterial growth and morphology" 3213: 3034: 2355: 2139: 2013: 1954: 1895: 1392: 1270: 1257: 1239: 1218: 1183: 1158: 1146: 1122: 829: 808:– climate-resistant spores that may form when environmental conditions become harsh 771: 730: 673: 543: 465: 434:. Understanding cyanobacterial morphogenesis is challenging, as there are numerous 235: 138: 5926: 5711:"Effects of tropical solar radiation on the motility of filamentous cyanobacteria" 5622: 4832:

Walsby, A. E. (1968). "Mucilage secretion and the movements of blue-green algae".

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Schirrmeister, Bettina E; Antonelli, Alexandre; Bagheri, Homayoun C (2011-02-14).

523: 347:

Earth, ranging from freshwater to marine and terrestrial habitats, including even

250:. Subsection III comprises multicellular, non-cell differentiating cyanobacteria ( 5673: 4817: 4800: 4421: 4374: 4357: 2509: 1525: 4614: 3940: 3578: 3163: 1452: 1308: 1274: 1214: 1178: 1079:, in order to provide the cells in the filament with nitrogen for biosynthesis. 1076: 1023: 1006: 834: 816: 812: 575: 491: 452: 396: 384: 300: 172: 122: 87: 5752:"Diel Migrations of Microorganisms within a Benthic, Hypersaline Mat Community" 5073: 4358:"Ultrafast photochemistry of Anabaena Sensory Rhodopsin: Experiment and theory" 4164: 3817: 2483:

Gumbart, James C.; Beeby, Morgan; Jensen, Grant J.; Roux, Benoît (2014-02-20).

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Whitton, Brian A.; Potts, Malcolm (2012). "Introduction to the Cyanobacteria".

1430: 5540:"A Model of Filamentous Cyanobacteria Leading to Reticulate Pattern Formation" 4801:"Secretion of the slime substance in Oscillatoria in relation to its movement" 4573: 4190: 3787: 3643: 3473: 3249: 3061: 3026: 2872: 1721: 1708:(2015). "Cytoskeletal crosstalk: when three different personalities team up". 1676: 1333: 1328: 1286: 1126: 1037: 602: 531: 484: 476: 435: 388: 360: 332: 271: 263: 255: 209: 199: 160:. The eukaryotic cytoskeleton is historically divided into three classes: the 102: 83: 6415: 4699: 4682: 4507: 4135: 3539: 3481: 3428: 3371: 3314: 3257: 3205: 3069: 2981: 2881: 2811: 2751: 2706: 2697: 2680: 2642: 2575: 2518: 2451: 2394: 2339: 2331: 2288: 2235: 2180: 2123: 2062: 1997: 1938: 1879: 1813: 1729: 1625: 1401: 133:, internal structures must dynamically adapt to the current requirements. In 5292: 3983: 3737: 2972: 2802: 2633: 2053: 1617: 871: 842: 653: 649: 565: 431: 348: 336: 306: 134: 6423: 6136: 6117: 6069: 5934: 5899: 5842: 5793: 5692: 5630: 5583: 5473: 5360: 5327:"Undirected motility of filamentous cyanobacteria produces reticulate mats" 5311: 5148: 5099: 5047: 5002: 4945: 4743: 4659: 4383: 4241: 4198: 4153: 4127: 3914: 3857: 3806: 3755: 3702: 3547: 3489: 3446: 3389: 3275: 3139: 3077: 2999: 2899: 2829: 2769: 2660: 2593: 2536: 2412: 2347: 2296: 2253: 2227: 2188: 2131: 1831: 1737: 1643: 1544: 1419: 6185: 6061: 6018: 5969: 5750:

Garcia-Pichel, Ferran; Mechling, Margaret; Castenholz, Richard W. (1994).

5191: 4902: 4735: 4342: 4293: 4079: 4071: 3363: 3332: 2469: 2080: 2005: 1946: 1887: 1429:

Modified material was copied from this source, which is available under a

1307:

Photographic negative projected onto a Petri dish containing a culture of

451:

Cyanobacteria present remarkable variability in terms of morphology: from

5139: 5122: 3187: 2743: 1804: 1327:

can position themselves quite precisely within their environment through

1324: 1316: 1222: 1134: 1098: 1063: 979: 921: 678: 642: 480: 439: 323: 275: 156: 75: 56: 5564: 5465: 4984: 4220:

Golden JW, Yoon HS (December 1998). "Heterocyst formation in Anabaena".

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5524: 4937: 4845: 4159: 3812: 3693: 3196: 1550: 1425: 1174: 1118: 974: 838: 805: 267: 215:

Cyanobacteria are today's only known prokaryotes capable of performing

191: 176: 168: 150: 5597:

Shaw, T.; Winston, M.; Rupp, C. J.; Klapper, I.; Stoodley, P. (2004).

5183: 5090: 5038: 5017: 4764:

Hansgirg A. (1883) "Bemerkungen über die Bewegungen der Oscillarien".

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Jasser, Iwona; Callieri, Cristiana (2017-02-11). "Picocyanobacteria".

2552:"Cyanobacterial Cell Walls: News from an Unusual Prokaryotic Envelope" 1836:

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forming long filaments of cells which can break into fragments called

2280: 2115: 1930: 1871: 1226: 1162: 1073: 410: 303: 274:). Whereas subsections III and IV form linear cell filaments (termed 146: 94: 5516: 3926: 3924: 208:

and even bacterial-specific cytoskeletal protein classes, including

5274: 3511:

Pinho, Mariana G.; Kjos, Morten; Veening, Jan-Willem (2013-08-16).

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is seen to be composed of one-cell-wide strands of connected cells.

1989: 1296: 1250: 1232: 1154: 1150: 1086: 851: 426:

Despite their morphological complexity, cyanobacteria contain all

290: 165: 29: 4967:

Duggan, Paula S.; Gottardello, Priscila; Adams, David G. (2007).

4449:. Developments in Sedimentology. Vol. 20. pp. 273–310. 3816:

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Ausmees, Nora; Kuhn, Jeffrey R; Jacobs-Wagner, Christine (2003).

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through pores on individual cells. Another theory suggests that

591: 195: 187: 6152:"Envelope structure of four gliding filamentous cyanobacteria" 5258:"Mechanisms for bacterial gliding motility on soft substrates" 3638:. Chichester, UK: John Wiley & Sons, Ltd. pp. 19–27. 2428:"Envelope structure of four gliding filamentous cyanobacteria" 982:

sheaths which can form tangles or mats, intermixed with other

6302:"A one-instant mechanism of phototaxis in the cyanobacterium 5072:

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Risser, Douglas D.; Chew, William G.; Meeks, John C. (2014).

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in an anaerobic environment due to its sensitivity to oxygen.

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Microphotographs of bundle-forming filamentous cyanobacteria

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Huber, Florian; Boire, Adeline; López, Magdalena Preciado;

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and self-organize into collective patterns and structures.

145:

polymers that assemble into stable or dynamic filaments or

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Many cyanobacteria form motile filaments of cells, called

6394:(1–3). Springer Science and Business Media LLC: 421–425. 1273:

is a genus of filamentous cyanobacterium named after the

4260:"Oxygen relations of nitrogen fixation in cyanobacteria" 4000:

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