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Genomic streamlining is a hypothesis in evolutionary biology and microbial ecology that suggests that there is a reproductive benefit to prokaryotes having a smaller genome size with less non-coding DNA and fewer non-essential genes.[1][2] There is a lot of variation in prokaryotic genome size, with the smallest free-living cell’s genome being roughly ten times smaller than the largest prokaryote.[3] Two of the bacterial taxa with the smallest genomes are Prochlorococcus and Pelagibacter ubique,[4][5] both highly abundant marine bacteria commonly found in oligotrophic regions. Similar reduced genomes have been found in uncultured marine bacteria, suggesting that genomic streamlining is a common feature of bacterioplankton[6]. This theory is typically used with reference to free-living organisms in oligotrophic environments.

Theory Overview

Comparison of genome sizes across select organisms

Genome streamlining theory states that certain prokaryotic genomes tend to be small in size in comparison to other prokaryotes, and all eukaryotes, due to selection against the retention of non-coding DNA. [2][1] The known advantages of small genome size include faster genome replication for cell division, fewer nutrient requirements, and easier co-regulation of multiple related genes, because gene density typically increases with decreased genome size[2]. This means that an organism with a smaller genome is likely to be more successful, or have higher fitness, than one hindered by excessive amounts of unnecessary DNA, leading to selection for smaller genome sizes.[2]

Some mechanisms that are thought to underlie genome streamlining include deletion bias and purifying selection.[1] Deletion bias is the phenomenon in bacterial genomes where the rate of DNA loss is naturally higher than the rate of DNA acquisition.[7][8] This is a passive process that simply results from the difference in these two rates.[8] Purifying selection is the process by which extraneous genes are selected against, making organisms lacking this genetic material more successful by effectively reducing their genome size.[7][9] Genes and non-coding DNA segments that are less crucial for an organism survival will be more likely to be lost over time.[9]

This selective pressure is stronger in large marine prokaryotic populations, because intra-species competition favours fast, efficient and inexpensive replication[2]. This is because large population sizes increase competition among members of the same species, and thus increases selective pressure and causes the reduction in genome size to occur more readily among organisms of large population sizes, like bacteria. [2] This may explain why genome streamlining seems to be particularly prevalent in prokaryotic organisms, as they tend to have larger population sizes than eukaryotes.[10]

It has also been proposed that having a smaller genome can help minimize overall cell size, which increases a prokaryotes surface-area to volume ratio.[11] A higher surface-area to volume ratio allows for more nutrient uptake proportional to their size, which allows them to outcompete other larger organisms for nutrients.[12][11] This phenomenon has been noted particularly in nutrient deplete waters.[11]

  1. ^ a b c Giovannoni, Stephen J.; Cameron Thrash, J.; Temperton, Ben (August 2014). "Implications of streamlining theory for microbial ecology". The ISME journal. 8 (8): 1553–1565. doi:10.1038/ismej.2014.60. ISSN 1751-7370. PMC 4817614. PMID 24739623.{{cite journal}}: CS1 maint: PMC format (link)
  2. ^ a b c d e f Sela, Itamar; Wolf, Yuri I.; Koonin, Eugene V. (2016-10-11). "Theory of prokaryotic genome evolution". Proceedings of the National Academy of Sciences. 113 (41): 11399–11407. doi:10.1073/pnas.1614083113.
  3. ^ Koonin, E. V.; Wolf, Y. I. (23 October 2008). "Genomics of bacteria and archaea: the emerging dynamic view of the prokaryotic world". Nucleic Acids Research. 36 (21): 6688–6719. doi:10.1093/nar/gkn668.
  4. ^ Giovannoni, S. J. (19 August 2005). "Genome Streamlining in a Cosmopolitan Oceanic Bacterium". Science. 309 (5738): 1242–1245. doi:10.1126/science.1114057.
  5. ^ Dufresne, A.; Salanoubat, M.; Partensky, F.; Artiguenave, F.; Axmann, I. M.; Barbe, V.; Duprat, S.; Galperin, M. Y.; Koonin, E. V.; Le Gall, F.; Makarova, K. S.; Ostrowski, M.; Oztas, S.; Robert, C.; Rogozin, I. B.; Scanlan, D. J.; de Marsac, N. T.; Weissenbach, J.; Wincker, P.; Wolf, Y. I.; Hess, W. R. (13 August 2003). "Genome sequence of the cyanobacterium Prochlorococcus marinus SS120, a nearly minimal oxyphototrophic genome". Proceedings of the National Academy of Sciences. 100 (17): 10020–10025. doi:10.1073/pnas.1733211100.
  6. ^ Swan, B. K.; Tupper, B.; Sczyrba, A.; Lauro, F. M.; Martinez-Garcia, M.; Gonzalez, J. M.; Luo, H.; Wright, J. J.; Landry, Z. C.; Hanson, N. W.; Thompson, B. P.; Poulton, N. J.; Schwientek, P.; Acinas, S. G.; Giovannoni, S. J.; Moran, M. A.; Hallam, S. J.; Cavicchioli, R.; Woyke, T.; Stepanauskas, R. (25 June 2013). "Prevalent genome streamlining and latitudinal divergence of planktonic bacteria in the surface ocean". Proceedings of the National Academy of Sciences. 110 (28): 11463–11468. doi:10.1073/pnas.1304246110.
  7. ^ a b Sela, Itamar; Wolf, Yuri I.; Koonin, Eugene V. (2016-10-11). "Theory of prokaryotic genome evolution". Proceedings of the National Academy of Sciences. 113 (41): 11399–11407. doi:10.1073/pnas.1614083113.
  8. ^ a b Mira, A.; Ochman, H.; Moran, N. A. (October 2001). "Deletional bias and the evolution of bacterial genomes". Trends in genetics: TIG. 17 (10): 589–596. ISSN 0168-9525. PMID 11585665.
  9. ^ a b Molina, Nacho; van Nimwegen, Erik (2008-1). "Universal patterns of purifying selection at noncoding positions in bacteria". Genome Research. 18 (1): 148–160. doi:10.1101/gr.6759507. ISSN 1088-9051. PMC 2134783. PMID 18032729. {{cite journal}}: Check date values in: |date= (help)CS1 maint: PMC format (link)
  10. ^ Lynch, Michael; Conery, John S. (2003-11-21). "The origins of genome complexity". Science (New York, N.Y.). 302 (5649): 1401–1404. doi:10.1126/science.1089370. ISSN 1095-9203. PMID 14631042.
  11. ^ a b c Chen, Bingzhang; Liu, Hongbin. "Relationships between phytoplankton growth and cell size in surface oceans: Interactive effects of temperature, nutrients, and grazing". Limnology and Oceanography. 55 (3): 965–972. doi:10.4319/lo.2010.55.3.0965.
  12. ^ Cotner, James; A. Biddanda, Bopaiah (2002-03-01). "Small Players, Large Role: Microbial Influence on Biogeochemical Processes in Pelagic Aquatic Ecosystems". Ecosystems. 5: 105–121. doi:10.1007/s10021-001-0059-3.