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Prochlorococcus
Genus of bacteria

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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Discovery

Although there had been several earlier records of very small chlorophyll-b-containing cyanobacteria in the ocean,78 Prochlorococcus was discovered in 19869 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.10 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 Prochlorococcus11 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.12

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.1314 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.15 Moreover, Prochlorococcus have adapted to use sulfolipids instead of phospholipids in their membranes to survive in phosphate deprived environments.16 This adaptation allows them to avoid competition with heterotrophs that are dependent on phosphate for survival.17 Typically, Prochlorococcus divide once a day in the subsurface layer or oligotrophic waters.18

Distribution

Prochlorococcus is abundant in the euphotic zone of the world's tropical oceans.19 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 individuals20 (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.21 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).22 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.23 Furthermore, Prochlorococcus are more plentiful in the presence of heterotrophs that have catalase abilities.24 Prochlorococcus do not have mechanisms to degrade reactive oxygen species and rely on heterotrophs to protect them.25 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.26

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.27 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.28

Genome

The genomes of several strains of Prochlorococcus have been sequenced.2930 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.31 Research has shown that a massive genome reduction occurred during the Neoproterozoic Snowball Earth, which was followed by population bottlenecks.32

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).33

DNA recombination, repair and replication

Marine Prochlorococcus cyanobacteria have several genes that function in DNA recombination, repair and replication. These include the recBCD gene complex whose product, exonuclease V, functions in recombinational repair of DNA, and the umuCD gene complex whose product, DNA polymerase V, functions in error-prone DNA replication.34 These cyanobacteria also have the gene lexA that regulates an SOS response system, probably a system like the well-studied E. coli SOS system that is employed in the response to DNA damage.35

Ecology

Ancestors of Prochlorococcus contributed to the production of early atmospheric oxygen.36 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.37 The size of Prochlorococcus (0.5 to 0.7 μm)38 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);39 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.40 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).41 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.424344 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.4546 It has been broken down by NCBI Taxonomy into two different subspecies, Low-light Adapted (LL) or High-light Adapted (HL).47 There are six clades within each subspecies.48

Low-light adapted

Prochlorococcus marinus subsp. marinus is associated with low-light adapted types.49 It is also further classified by sub-ecotypes LLI-LLVII, where LLII/III has not been yet phylogenetically uncoupled.5051 LV species are found in highly iron scarce locations around the equator, and as a result, have lost several ferric proteins.52 The low-light adapted subspecies is otherwise known to have a higher ratio of chlorophyll b2 to chlorophyll a2,53 which aids in its ability to absorb blue light.54 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.5556 Their genomes can range from 1,650,000 to 2,600,000 basepairs in size.57

High-light adapted

Prochlorococcus marinus subsp. pastoris is associated with high-light adapted types.58 It can be further classified by sub-ecotypes HLI-HLVI.5960 HLIII, like LV, is also located in an iron-limited environment near the equator, with similar ferric adaptations.61 The high-light adapted subspecies is otherwise known to have a low ratio of chlorophyll b2 to chlorophyll a2.62 High-light adapted strains inhabit depths between 25 and 100 m.63 Their genomes can range from 1,640,000 to 1,800,000 basepairs in size.64

Metabolism

Most cyanobacterium are known to have an incomplete tricarboxylic acid cycle (TCA).6566 In this process, 2-oxoglutarate decarboxylase (2OGDC) and succinic semialdehyde dehydrogenase (SSADH), replace the enzyme 2-oxoglutarate dehydrogenase (2-OGDH).67 Normally, when this enzyme complex joins with NADP+, it can be converted to succinate from 2-oxoglutarate (2-OG).68 This pathway is non-functional in Prochlorococcus,69 as succinate dehydrogenase has been lost evolutionarily to conserve energy that may have otherwise been lost to phosphate metabolism.70

Strains

StrainSubtypeSource
MIT9515HLI71
EQPAC1HLI72
MED4HLI73
XMU1401HLII74
MIT0604HLII75
AS9601HLII76
GP2HLII77
MIT9107HLII78
MIT9116HLII79
MIT9123HLII80
MIT9201HLII81
MIT9202HLII82
MIT9215HLII83
MIT9301HLII84
MIT9302HLII85
MIT9311HLII86
MIT9312HLII87
MIT9314HLII88
MIT9321HLII89
MIT9322HLII90
MIT9401HLII91
SBHLII92
XMU1403LLI93
XMU1408LLI94
MIT0801LLI95
NATL1ALLI96
NATL2ALLI97
PAC1LLI98
LGLLII/III99
MIT0601LLII/III100
MIT0602LLII/III101
MIT0603LLII/III102
MIT9211LLII/III103
SS35LLII/III104
SS52LLII/III105
SS120LLII/III106
SS2LLII/III107
SS51LLII/III108
MIT0701LLIV109
MIT0702LLIV110
MIT0703LLIV111
MIT9303LLIV112
MIT9313LLIV113
MIT1303LLIV114
MIT1306LLIV115
MIT1312LLIV116
MIT1313LLIV117
MIT1318LLIV118
MIT1320LLIV119
MIT1323LLIV120
MIT1327LLIV121
MIT1342LLIV122

Table modified from 123

See also

Further reading

References

  1. Munn, C. (2011). Marine Microbiology: Ecology and applications (2nd ed.). Garland Science.[page needed] /wiki/Wikipedia:Citing_sources ↩

  2. Chimileski, Scott; Kolter, Roberto (25 September 2017). Life at the Edge of Sight. Belknap Press. ISBN 978-0-674-97591-0. Retrieved 2018-01-26.[page needed] 978-0-674-97591-0 ↩

  3. Tolonen AC, Aach J, Lindell D, Johnson ZI, Rector T, Steen R, Church GM, Chisholm SW (October 2006). "Global gene expression of Prochlorococcus ecotypes in response to changes in nitrogen availability". Molecular Systems Biology. 2 (1). 53. doi:10.1038/msb4100087. PMC 1682016. PMID 17016519. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1682016 ↩

  4. 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. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2151091 ↩

  5. Munn, C. (2011). Marine Microbiology: Ecology and applications (2nd ed.). Garland Science.[page needed] /wiki/Wikipedia:Citing_sources ↩

  6. 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. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2151091 ↩

  7. Johnson, P.W.; Sieburth, J.M. (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. /wiki/John_McNeill_Sieburth ↩

  8. Gieskes, W.W.C.; Kraay, G.W. (1983). "Unknown chlorophyll a derivatives in the North Sea and the tropical Atlantic Ocean revealed by HPLC analysis". Limnology and Oceanography. 28 (4): 757–766. Bibcode:1983LimOc..28..757G. doi:10.4319/lo.1983.28.4.0757. https://doi.org/10.4319%2Flo.1983.28.4.0757 ↩

  9. Chisholm, S.W.; Olson, R.J.; Zettler, E.R.; Waterbury, J.; Goericke, R.; Welschmeyer, N. (1988). "A novel free-living prochlorophyte occurs at high cell concentrations in the oceanic euphotic zone". Nature. 334 (6180): 340–3. Bibcode:1988Natur.334..340C. doi:10.1038/334340a0. S2CID 4373102. /wiki/Nature_(journal) ↩

  10. "The Crafoord Prize in Biosciences 2019". Royal Swedish Academy of Sciences. January 17, 2019. Retrieved April 26, 2022. https://www.crafoordprize.se/press_release/the-crafoord-prize-in-biosciences-2019 ↩

  11. Chisholm, S.W.; Frankel, S.L.; Goericke, R.; Olson, R.J.; Palenik, B.; Waterbury, J.B.; West-Johnsrud, L.; Zettler, E.R. (1992). "Prochlorococcus marinus nov. gen. nov. sp.: an oxyphototrophic marine prokaryote containing divinyl chlorophyll a and b". Archives of Microbiology. 157 (3): 297–300. Bibcode:1992ArMic.157..297C. doi:10.1007/BF00245165. S2CID 32682912. /wiki/Sallie_W._Chisholm ↩

  12. "Prochlorococcus marinus". NCBI. Retrieved 2022-04-25. https://www.ncbi.nlm.nih.gov/data-hub/taxonomy/1219/ ↩

  13. Biller, Steven J.; Berube, Paul M.; Lindell, Debbie; Chisholm, Sallie W. (1 December 2014). "Prochlorococcus: the structure and function of collective diversity" (PDF). Nature Reviews Microbiology. 13 (1): 13–27. doi:10.1038/nrmicro3378. hdl:1721.1/97151. PMID 25435307. S2CID 18963108. /wiki/Debbie_Lindell ↩

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  15. Partensky F, Hess WR, Vaulot D (1999). "Prochlorococcus, a marine photosynthetic prokaryote of global significance". Microbiology and Molecular Biology Reviews. 63 (1): 106–127. doi:10.1128/MMBR.63.1.106-127.1999. PMC 98958. PMID 10066832. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC98958 ↩

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  19. Chisholm, S.W.; Frankel, S.; Goericke, R.; Olson, R.; Palenik, B.; Waterbury, J.; West-Johnsrud, L.; Zettler, E. (1992). "Prochlorococcus marinus nov. gen. nov. sp.: an oxyphototrophic marine prokaryote containing divinyl chlorophyll a and b.". Archives of Microbiology. 157 (3): 297–300. Bibcode:1992ArMic.157..297C. doi:10.1007/bf00245165. S2CID 32682912. /wiki/Bibcode_(identifier) ↩

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