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Purple bacteria grown in Winogradsky column

Purple bacteria are gram-negative photosynthetic prokaryotes, belonging to the phylum of Proteobacteria with a characteristic purple color given by the presence of carotenoids, accessory pigments responsible for capturing light and photoprotection. Purple bacteria are divided into purple sulfur bacteria (Chromatiales), species of gamma proteobacteria and, and purple non-sulfur bacteria (Rhodospirillaceae), species of alpha and beta Proteobacteria [1]. Purple bacteria are anoxygenic phototrophs widely spread in nature, but especially in aquatic environments, where there are anoxic conditions that favor the synthesis of their pigments [2]. Purple bacteria use a type II photosystem and have type a or b bacteriochlorophylls.

Taxonomy

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Purple bacteria belong to phylum of Proteobacteria. This phylum was established by Carl Woese in 1987 calling it “purple bacteria and their relatives” even if this is not appropriate because most of them are not purple or photosynthetic[3]. Phylum Proteobacteria is divided into six classes:Alphaproteobacteria, Betaproteobacteria, Gammaproteobacteria, Deltaproteobacteria, Epsilonproteobacteria and Zetaproteobcateria. Purple bacteria are distributed between 3 classes:Alphaproteobacteria, Betaproteobacteria, Gammaproteobacteria.[4] each caracterized by a photosynthetic phenotype.

Alpha subdivision contains different photosynthetic purple bacteria species (for instance: Rhodospirillum, Rhodopseudomonas and Rhodomicrobium) but include also some non-photosyntetic purple ones of genera with nitrogen metabolism ( Rhizobium , Nitrobacter) whereas in betaproteobacteria subdivision there are few photosynthetic species. Gammaproteobacteria class is divided into 3 subgroups : gamma-1, gamma-2, gamma-3. In gamma-1 subgroup there are the purple photosynthetic bacteria that produce molecular sulfur (Chromatiaceae group and Ectothiorhodospiraceae group) and also the non-photosyntetic species ( as Nitrosococcus oceani[5])

Purple sulfur bacteria and purple nunsulfur bacteria were distinguished on the basis of physiological factors of their tolerance and utilization of sulfide: was considered that purple sulfur bacteria tolerate millimolar levels of sulfide and oxidized sulfide to sulfur globules stored intracellulary while purple nonsulfur bacteria species did neither.[6] This kind of classification was not absoluted. It’s was refuted with classic chemostat experiments by Hansen and Van Gemerden (1972) that demonstrate the growing of many purple nonsulfur bacteria species at low levels of sulfide (0.5mM) and in so doing , oxidize sulfide to S0, S4O62-, or SO42-. The important distinction that remains from these two different metabolisms is that: any S0 formed by purple nonsulfur bacteria is not stored intracellularly but is deposited outside the cell[7] (even if there are exception for this as Ectothiorhodospiraceae). So if grown on sulfide it is easy to differentiate purple sulfur bacteria from purple non sulfur bacteria because the microscopically globules of S0 are formed.[8]

Biogeochemical cycles

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Purple bacteria are involved in the biogeochemical cycles of different nutrients. In fact they are able to photoautotrophically fix carbon, or to consume it photoheterotrophically; in both cases in anoxic conditions. However the most important role is played by consuming hydrogen sulphide: a highly toxic substance for plants, animals and other bacteria. In fact, the oxidation of hydrogen sulphide by purple bacteria produces non-toxic forms of sulfur, such as elemental sulfur and sulphate[9]. In addition, almost all non-sulfur purple bacteria are able to fix nitrogen (N2+8H+--->2NH3+H2)[10], and Rba Sphaeroides, an alpha proteobacter, is capable of reducing nitrate to molecular nitrogen by denitrification[11].

Metabolism

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Purple Bacteria are able to perform different metabolisms that allow them to adapt to different and even exteme enviromental conditions. They are mainly photoautotrophs carrying out anoxygenic photosyntesis, but they are also known to be chemoautotrophs or photoheterotrophs. Since pigments synthesis does not take place in presence of oxygen, phototrophic growth only occurs in anoxic and light conditions[12]. Howerver purple bacteria can also grow in dark and oxic enviroments. Infact they can be mixotrophs, capable of aerobic respiration or fermentation[13] basing on the concentration of oxygen and availability of light[14].

Photosynthesis

[edit]

Photosynthetic Unit

Purple Bacteria use bacteriochlorophyll and carotenoids to obtain the light energy for the photosynthesis. Electron transfer and photosynthetic reactions occur at the cell membrane in the photosynthetic unit which is composed by the light-harvesting complexes LHI and LHII and the photosynthetic reaction centre where the charge separation reaction occurs.[15] These structures are located in the intracytoplasmic membrane, areas of the cytoplasmic membrane invaginated to form vesicle sacs, tubules, or single-paired or stacked lamellar sheets [16]. Light-harvesting complexes are involved in the energy transfer to the reaction centre. These are integral membrane protein complexes consisting of monomers of α- and β- apoproteins, each one binding molecules of bacteriochlorophyll and carotenoids non-covalently. LHI is directly associated with the reaction centre forming a polymeric ring-like structure around it. LHI has a absorption maximum at 875 nm and it contains most of the bacteriochlorophyll of the photosynthetic unit. LHII contains less bacteriochlorophylls, has lower absorption maximum (850 nm) and is not present in all purple bacteria[17]. Moreover, the photosynthetic unit in Purple Bacteria shows great plasticity, being able to adapt to the constantly changing light conditions. Infact these microorganisms are able to rearrange the composition and the concentration of the pigments, and consequently the absorption spectrum, in response to light variation. [18]

Electron donors and carbon sources for anabolism

Purple bacteria also transfer electrons from external electron donors directly to cytochrome bc1 to generate NADH or NADPH used for anabolism.[19]. They are anoxygenic because they do not use water as an electron donor to produce oxygen. Purple sulfur bacteria (PSB), use sulfide; sulfur; thiosulfate; or hydrogen as electron donors.[20] In addition, some species use ferrous iron as electron donor [21] and one strain of Thiocapsa can use nitrite. Finally, even if the purple sulfur bacteria are typically photoautotroph, some of them are photoheterotroph and use differents carbon sources and electron donor such as organic acids and fatty acids. On the other hand purple non-sulfur bacteria, typically use hydrogen as an electron donor, but can also use sulfide at lower concentrations compared to PSB and some species can use thiosulfate or ferrous iron as electron donor [22]. In contrast to the purple sulfur bacteria, the purple non sulfur bacteria are mostly photoheterotrophic and can use as elctron donor and carbon sources such as sugars, amino acids, organic acids, and aromatic compounds like toluene or benzoate.


Distribution

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Purple bacteria inhabit illuminated anoxic aquatic and terrestrial environments. Even if sometimes the two major groups of purple bacteria, purple sulfur bacteria and purple nonsulfur bacteria, coexist in the same habitat, they occupy different niches. Purple sulfur bacteria are strongly photoautotrophs and are not adapted to an efficient metabolism and growth in the dark. A different speech applies to purple nonsulfur bacteria that are strongly photoheterotrophs, even if they are capable of photoautotrophy, and are equipped for living in dark environments. Purple sulfur bacteria can be found in different ecosystems with enough sulfate and light, some examples are shallow lagoons polluted by sewage or deep waters of lakes, in which they could even bloom. Blooms can both involve a single or a mixture of species. They can also be found in microbial mats where the lower layer decomposes and sulfate-reduction occurs[23].

Purple non sulfur bacteria can be found in both illuminated and dark environments with lack of sulfide. However, they hardly form blooms with sufficiently high concentration to be visible without enrichment techniques[24].

Purple bacteria have evolved effective strategies for photosynthesis in extreme environments, in fact they are quite successful in harsh habitats. In the 1960s the first halophiles and acidophiles of the genus Ectothiorhodospira were discovered. In the 1980s Thermochromatium tepidum, a thermophilic purple bacterium that can be found in North American hot springs, was isolated for the first time[25].

Ecology

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  • Ecological niches

Quantity and quality of light

Several studies have shown that a strong accumulation of phototrophic sulfur bacteria has been observed between 2 and 20 meters deep (in some cases even 30 m) of pelagic environments[26]. This is due to the fact that in some environments the light transmission for various populations of phototrophic sulfur bacteria varies with a density from 0.015 to 10%[27]. Furthermore, Chromatiaceae have been found in chemocline environments over ≤20 m depths. The correlation between anoxygenic photosynthesis and the availability of solar radiation suggests that light is the main factor controlling all the activities of phototrophic sulfur bacteria. The density of pelagic communities of phototrophic sulfur bacteria extends beyond a depth range of 10 cm [28], while the less dense population (found in the Black Sea (0.068–0.94 μg BChle/dm-3), is scattered over an interval of 30 m[29]. Communities of phototrophic sulfur bacteria located in the coastal sediments of sandy, saline or muddy beaches live in an environment with a higher light gradient , limiting growth to the highest value between 1.5-5mm of the sediments[30] . At the same time, biomass densities of 900 mg bacteriochlorophyll/dm-3 can be attained in these latter systems[31].

Temperature and salinity

Purple sulfur bacteria (such as Green sulfur bacteria) typically form blooms in non-thermal aquatic ecosystems, some members have been found in hot springs[32] . For example Chlorobaculum tepidum can only be found in some hot springs in New Zealand at a ph value between 4.3 and 6.2 and at a temperature above 56 ° C. Another example are Thermochromatium tepidum, has been found in several hot springs in western North America at temperatures above 58 ° C and may represent the most thermophilic proteobacteria existent[33]. Of the purple sulfur bacteria, many members of the Chromatiaceae family are often found in fresh water and marine environments. About 10 species of Chromatiaceae are halophilic [34].

Syntrophy and symbioses

Like green sulfur bacteria, purple sulfur bacteria are also capable of symbiosis. In fact, it was noted that they rapidly create stable associations[35] between other purple sulfur bacteria and sulfur- or sulfate-reducing bacteria. These associations are based on a cycle of sulfur but not carbon compounds. Thus, a simultaneous growth of two bacteria partners takes place, which are fed by the oxidation of organic carbon and light substrates. Experiments with Chromatiaceae have pointed out that cell aggregates consisting of sulfate-reducing Proteobacterium Desulfocapsa Thiozymogenes and small cells of Chromatiaceae have been observed in the chemocline of an alpine mermocitic lake[36] .

It was also noted that the purple sulfur bacteria Chromatium weissei often establish commensal associations with epibiotic bacteria[37]. An unidentified epibiont attaches to healthy Chromatium cells by performing a lysis process on host cells, a methodology very similar to that of Vampirococcus[38]. It seems that the epibiont grows chemotrophically on carbon compounds excreted by the purple sulfur bacteria.


References

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  1. ^ Imhoff, Johannes F.; Hiraishi, Akira; Süling, Jörg (2015), "Anoxygenic Phototrophic Purple Bacteria", Bergey's Manual of Systematics of Archaea and Bacteria, American Cancer Society, pp. 1–23, doi:10.1002/9781118960608.bm00002, ISBN 978-1-118-96060-8, retrieved 2020-11-21
  2. ^ Cohen-Bazire, Germaine; Sistrom, W. R.; Stanier, R. Y. (1957). "Kinetic studies of pigment synthesis by non-sulfur purple bacteria". Journal of Cellular and Comparative Physiology. 49 (1): 25–68. doi:10.1002/jcp.1030490104. ISSN 1553-0809. PMID 13416343.
  3. ^ Stackebrandt, E.; Murray, R. G. E.; Truper, H. G. (1 July 1988). "Proteobacteria classis nov., a Name for the Phylogenetic Taxon That Includes the "Purple Bacteria and Their Relatives"". International Journal of Systematic Bacteriology. 38 (3): 321–325. doi:10.1099/00207713-38-3-321. ISSN 0020-7713.
  4. ^ Takaichi, Shinichi (2009). "Distribution and Biosynthesis of Carotenoids". The Purple Phototrophic Bacteria. Advances in Photosynthesis and Respiration. Vol. 28. pp. 97–117. doi:10.1007/978-1-4020-8815-5_6. ISBN 978-1-4020-8814-8.
  5. ^ Woese, C.R.; Weisburg, W.G.; Hahn, C.M.; Paster, B.J.; Zablen, L.B.; Lewis, B.J.; Macke, T.J.; Ludwig, W.; Stackebrandt, E. (June 1985). "The Phylogeny of Purple Bacteria: The Gamma Subdivision". Systematic and Applied Microbiology. 6 (1): 25–33. Bibcode:1985SyApM...6...25W. doi:10.1016/S0723-2020(85)80007-2.
  6. ^ Niel, C. B. (1932). "On the morphology and physiology of the purple and green sulphur bacteria". Archiv for Mikrobiologie. 3 (1): 1–112. Bibcode:1932ArMic...3....1V. doi:10.1007/BF00454965. S2CID 19597530.
  7. ^ Hansen, Theo A.; Gemerden, Hans (1972). "Sulfide utilization by purple nonsulfur bacteria". Archiv for Mikrobiologie. 86 (1): 49–56. Bibcode:1972ArMic..86...49H. doi:10.1007/BF00412399. PMID 4628180. S2CID 7410927.
  8. ^ Madigan, Michael T.; Jung, Deborah O. (2009). "An Overview of Purple Bacteria: Systematics, Physiology, and Habitats". The Purple Phototrophic Bacteria. Advances in Photosynthesis and Respiration. Vol. 28. pp. 1–15. doi:10.1007/978-1-4020-8815-5_1. ISBN 978-1-4020-8814-8.
  9. ^ Madigan, Michael T.; Jung, Deborah O. (2009), Hunter, C. Neil; Daldal, Fevzi; Thurnauer, Marion C.; Beatty, J. Thomas (eds.), "An Overview of Purple Bacteria: Systematics, Physiology, and Habitats", The Purple Phototrophic Bacteria, vol. 28, Dordrecht: Springer Netherlands, pp. 1–15, doi:10.1007/978-1-4020-8815-5_1, ISBN 978-1-4020-8814-8, retrieved 2020-11-21
  10. ^ Madigan, Michael T. (1995), Blankenship, Robert E.; Madigan, Michael T.; Bauer, Carl E. (eds.), "Microbiology of Nitrogen Fixation by Anoxygenic Photosynthetic Bacteria", Anoxygenic Photosynthetic Bacteria, Advances in Photosynthesis and Respiration, vol. 2, Dordrecht: Springer Netherlands, pp. 915–928, doi:10.1007/0-306-47954-0_42, ISBN 978-0-306-47954-0, retrieved 2020-12-10
  11. ^ Satoh, Toshio; Hoshino, Yasuo; Kitamura, Hiroshi (1976-07-01). "Rhodopseudomonas sphaeroides forma sp. denitrificans, a denitrifying strain as a subspecies of Rhodopseudomonas sphaeroides". Archives of Microbiology. 108 (3): 265–269. Bibcode:1976ArMic.108..265S. doi:10.1007/BF00454851. ISSN 1432-072X. PMID 1085137. S2CID 20375188.
  12. ^ Keppen, O. I.; Krasil’nikova, E. N.; Lebedeva, N. V.; Ivanovskii, R. N. (2013-09-01). "Comparative study of metabolism of the purple photosynthetic bacteria grown in the light and in the dark under anaerobic and aerobic conditions". Microbiology. 82 (5): 547–553. doi:10.1134/S0026261713050056. ISSN 1608-3237. PMID 25509391. S2CID 254842108.
  13. ^ Tsygankov, A. A.; Khusnutdinova, A. N. (2015-01-01). "Hydrogen in metabolism of purple bacteria and prospects of practical application". Microbiology. 84 (1): 1–22. doi:10.1134/S0026261715010154. ISSN 1608-3237. S2CID 254835335.
  14. ^ Hädicke, Oliver; Grammel, Hartmut; Klamt, Steffen (2011-09-25). "Metabolic network modeling of redox balancing and biohydrogen production in purple nonsulfur bacteria". BMC Systems Biology. 5 (1): 150. doi:10.1186/1752-0509-5-150. ISSN 1752-0509. PMC 3203349. PMID 21943387.
  15. ^ Thorsten Ritz; Ana Damjanovic; Klaus Schulten (2002). "The Quantum Physics of Photosynthesis". ChemPhysChem. 3 (3): 243–248. doi:10.1002/1439-7641(20020315)3:3<243::AID-CPHC243>3.0.CO;2-Y. PMID 12503169. Retrieved 15 March 2002.
  16. ^ "Structure, Function and Formation of Bacterial Intracytoplasmic Membranes". ResearchGate. Retrieved 2017-10-08.
  17. ^ Francke, Christof; Amesz, Jan (1995-11-01). "The size of the photosynthetic unit in purple bacteria". Photosynthesis Research. 46 (1): 347–352. Bibcode:1995PhoRe..46..347F. doi:10.1007/BF00020450. ISSN 1573-5079. PMID 24301602. S2CID 23254767.
  18. ^ Brotosudarmo, Tatas Hardo Panintingjati; Limantara, Leenawaty; Heriyanto; Prihastyanti, Monika Nur Utami (2015-01-01). "Adaptation of the Photosynthetic Unit of Purple Bacteria to Changes of Light Illumination Intensities". Procedia Chemistry. 2nd Humboldt Kolleg in conjunction with International Conference on Natural Sciences 2014, HK-ICONS 2014. 14: 414–421. doi:10.1016/j.proche.2015.03.056. ISSN 1876-6196.
  19. ^ "The architecture and function of the light-harvesting apparatus of purple bacteria: from single molecules to in vivo membranes". ProQuest. Retrieved 2017-10-08.
  20. ^ Basak, Nitai; Das, Debabrata (2007-01-01). "The Prospect of Purple Non-Sulfur (PNS) Photosynthetic Bacteria for Hydrogen Production: The Present State of the Art". World Journal of Microbiology and Biotechnology. 23 (1): 31–42. doi:10.1007/s11274-006-9190-9. ISSN 0959-3993. S2CID 84224465.
  21. ^ Ehrenreich, A.; Widdel, F. (1994-12-01). "Anaerobic oxidation of ferrous iron by purple bacteria, a new type of phototrophic metabolism". Applied and Environmental Microbiology. 60 (12): 4517–4526. Bibcode:1994ApEnM..60.4517E. doi:10.1128/aem.60.12.4517-4526.1994. ISSN 0099-2240. PMC 202013. PMID 7811087.
  22. ^ Brune, Daniel C. (1995), Blankenship, Robert E.; Madigan, Michael T.; Bauer, Carl E. (eds.), "Sulfur Compounds as Photosynthetic Electron Donors", Anoxygenic Photosynthetic Bacteria, Advances in Photosynthesis and Respiration, vol. 2, Dordrecht: Springer Netherlands, pp. 847–870, doi:10.1007/0-306-47954-0_39, ISBN 978-0-306-47954-0, retrieved 2020-12-10
  23. ^ Madigan, Michael T.; Jung, Deborah O. (2009). "An Overview of Purple Bacteria: Systematics, Physiology, and Habitats". The Purple Phototrophic Bacteria. Advances in Photosynthesis and Respiration. Vol. 28. pp. 1–15. doi:10.1007/978-1-4020-8815-5_1. ISBN 978-1-4020-8814-8.
  24. ^ Siefert, E.; Irgens, R. L.; Pfennig, N. (1 January 1978). "Phototrophic purple and green bacteria in a sewage treatment plant". Applied and Environmental Microbiology. 35 (1): 38–44. Bibcode:1978ApEnM..35...38S. doi:10.1128/aem.35.1.38-44.1978. ISSN 0099-2240. PMID 623470. S2CID 28567941.
  25. ^ Imhoff, Johannes F. (2017). "Anoxygenic Phototrophic Bacteria from Extreme Environments". Modern Topics in the Phototrophic Prokaryotes. Springer International Publishing. pp. 427–480. doi:10.1007/978-3-319-46261-5_13. ISBN 978-3-319-46259-2. {{cite book}}: |journal= ignored (help)
  26. ^ Herbert, Rodney A.; Ranchou-Peyruse, Anthony; Duran, Robert; Guyoneaud, Rémy; Schwabe, Stephanie (2005). "Characterization of purple sulfur bacteria from the South Andros Black Hole cave system: highlights taxonomic problems for ecological studies among the genera Allochromatium and Thiocapsa". Environmental Microbiology. 7 (8): 1260–1268. Bibcode:2005EnvMi...7.1260H. doi:10.1111/j.1462-2920.2005.00815.x. ISSN 1462-2920. PMID 16011763.
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  29. ^ Manske, Ann K.; Glaeser, Jens; Kuypers, Marcel M. M.; Overmann, Jörg (2005). "Physiology and Phylogeny of Green Sulfur Bacteria Forming a Monospecific Phototrophic Assemblage at a Depth of 100 Meters in the Black Sea". Applied and Environmental Microbiology. 71 (12): 8049–8060. Bibcode:2005ApEnM..71.8049M. doi:10.1128/aem.71.12.8049-8060.2005. PMC 1317439. PMID 16332785.
  30. ^ Van Gemerden, Hans; Mas, Jordi (1995), "Ecology of Phototrophic Sulfur Bacteria", Advances in Photosynthesis and Respiration, Dordrecht: Springer Netherlands, pp. 49–85, doi:10.1007/0-306-47954-0_4, ISBN 978-0-7923-3681-5, retrieved 2020-11-22
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  36. ^ Tonolla, Mauro; Demarta, Antonella; Peduzzi, Sandro; Hahn, Dittmar; Peduzzi, Raffaele (2000-02-01). "In Situ Analysis of Sulfate-Reducing Bacteria Related to Desulfocapsa thiozymogenes in the Chemocline of Meromictic Lake Cadagno (Switzerland)". Applied and Environmental Microbiology. 66 (2): 820–824. Bibcode:2000ApEnM..66..820T. doi:10.1128/AEM.66.2.820-824.2000. ISSN 1098-5336. PMC 91902. PMID 10653757.
  37. ^ Clarke, K. J.; Finlay, B. J.; Vicente, E.; Lloréns, H.; Miracle, M. R. (1993-06-01). "The complex life-cycle of a polymorphic prokaryote epibiont of the photosynthetic bacterium Chromatium weissei". Archives of Microbiology. 159 (6): 498–505. Bibcode:1993ArMic.159..498C. doi:10.1007/BF00249026. ISSN 1432-072X. S2CID 30475947.
  38. ^ Overmann, Jörg (2008), Hell, Rüdiger; Dahl, Christiane; Knaff, David; Leustek, Thomas (eds.), "Ecology of Phototrophic Sulfur Bacteria", Sulfur Metabolism in Phototrophic Organisms, Advances in Photosynthesis and Respiration, vol. 27, Dordrecht: Springer Netherlands, pp. 375–396, doi:10.1007/978-1-4020-6863-8_19, ISBN 978-1-4020-6862-1, retrieved 2020-11-22
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