symbiogenesis

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Electron micrograph of a mitochondrion showing its mitochondrial matrix and membranes

Symbiogenesis, or endosymbiotic theory, is an evolutionary theory which explains the origin of eukaryotic cells from prokaryotes. It states that several key organelles of eukaryotes originated as symbiosis between separate single-celled organisms. According to this theory, mitochondria and plastids (e.g. chloroplasts), and possibly other organelles, represent formerly free-living bacteria that were taken inside another cell as an endosymbiont, around 1.5 billion years ago. Molecular and biochemical evidence suggest that the mitochondrion developed from proteobacteria (in particular, Rickettsiales, the SAR11 clade,^[1]^[2] or close relatives) and the chloroplast from cyanobacteria (in particular, N₂-fixing filamentous cyanobacteria^[3]^[4]).

History

Endosymbiotic theory

The endosymbiotic (Greek: ἔνδον endon "within", σύν syn "together" and βίωσις biosis "living") theories were first articulated by the Russian botanist Konstantin Mereschkowski in 1910,^[5] although the fundamental elements of the theory were described in a paper five years earlier.^[6]^[7] Mereschkowski was familiar with work by botanist Andreas Schimper, who had observed in 1883 that the division of chloroplasts in green plants closely resembled that of free-living cyanobacteria, and who had himself tentatively proposed (in a footnote) that green plants had arisen from a symbiotic union of two organisms.^[8] Ivan Wallin extended the idea of an endosymbiotic origin to mitochondria in the 1920s.^[9]^[10] A Russian botanist Boris Kozo-Polyansky was the first to explain the theory in terms of Darwinian evolution.^[11] In his 1924 book Symbiogenesis: A New Principle of Evolution he wrote, "The theory of symbiogenesis is a theory of selection relying on the phenomenon of symbiosis."^[12] These theories were initially dismissed or ignored. More detailed electron microscopic comparisons between cyanobacteria and chloroplasts (for example studies by Hans Ris published in 1961^[13]), combined with the discovery that plastids and mitochondria contain their own DNA^[14] (which by that stage was recognized to be the hereditary material of organisms) led to a resurrection of the idea in the 1960s.

The endosymbiotic theory was advanced and substantiated with microbiological evidence by Lynn Margulis in a 1967 paper, On the origin of mitosing cells.^[15] In her 1981 work Symbiosis in Cell Evolution she argued that eukaryotic cells originated as communities of interacting entities, including endosymbiotic spirochaetes that developed into eukaryotic flagella and cilia. This last idea has not received much acceptance, because flagella lack DNA and do not show ultrastructural similarities to bacteria or archaea (see also: Evolution of flagella and Prokaryotic cytoskeleton). According to Margulis and Dorion Sagan,^[16] "Life did not take over the globe by combat, but by networking" (i.e., by cooperation). The possibility that peroxisomes may have an endosymbiotic origin has also been considered, although they lack DNA. Christian de Duve proposed that they may have been the first endosymbionts, allowing cells to withstand growing amounts of free molecular oxygen in the Earth's atmosphere. However, it now appears that they may be formed de novo, contradicting the idea that they have a symbiotic origin.^[17]

It is thought that over millennia these endosymbionts transferred some of their own DNA to the host cell's nucleus (called "endosymbiotic gene transfer") during the evolutionary transition from a symbiotic community to an instituted eukaryotic cell. The endosymbiotic theory is considered to be a type of saltational evolution.^[18]

One model for the origin of mitochondria and plastids.

From endosymbionts to organelles

According to Keeling and Archibald,^[19] the usual way to distinguish organelles from endosymbionts is by their reduced genome sizes. As an endosymbiont evolves into an organelle, most of their genes are transferred to the host cell genome. The host cell and organelle need to develop a transport mechanism that enables transfer back of the protein products needed by the organelle but now manufactured by the cell. However, using the example of the freshwater amoeboid Paulinella chromatophora, which contains chromatophores found to be evolved from cyanobacteria, these authors argue that this is not the only possible criterion, another one being that the host cell has assumed control of the regulation of the former endosymbiont's division, bringing it in synchrony with the cell's own division.^[19] Nowack and her colleagues^[20] performed gene sequencing on the chromatophore (1.02Mb) and found that only 867 proteins were encoded by these photosynthetic cells. Comparisons with their closest free living cyanobacteria of the genus Synechococcus (having a genome size of 3Mb with 3300 genes) revealed that chromatophores underwent a drastic genome shrinkage. Chromatophores contained genes that were accountable for photosynthesis but were deficient in genes that could carry out other biosynthetic functions signifying that these endosymbiotic cells were highly dependent on their hosts for their survival and growth mechanisms. Thus, these chromatophores were found to be non-functional for organelle-specific purposes when compared to mitochondria and plastids. This distinction could have promoted the early evolution of photosynthetic organelles.

Evidence

Evidence that mitochondria and plastids arose from bacteria is as follows:^[21]^[22]^[23]

New mitochondria and plastids are formed only through a process similar to binary fission.
If a cell's mitochondria or chloroplasts are removed, they do not have the means to create new ones.^[24] For example, in some algae, such as Euglena, the plastids can be destroyed by certain chemicals or prolonged absence of light without otherwise affecting the cell. In such a case, the plastids will not regenerate.
Transport proteins called porins are found in the outer membranes of mitochondria and chloroplasts, are also found in bacterial cell membrane.^[25]^[26]^[27]
A membrane lipid cardiolipin is exclusively found in the inner mitochondrial membrane and bacterial cell membrane.^[28]
Both mitochondria and plastids contain single circular DNA that is different from that of the cell nucleus and that is similar to that of bacteria (both in their size and structure).
The genomes, including the specific genes, are basically similar between mitochondria and the Rickettsial bacteria.^[29]
Genome comparisons indicate that cyanobacteria contributed to the genetic origin of plastids.^[30]
DNA sequence analysis and phylogenetic estimates suggest that nuclear DNA contains genes that probably came from plastids.
These organelles' ribosomes are like those found in bacteria (70S).
Proteins of organelle origin, like those of bacteria, use N-formylmethionine as the initiating amino acid.
Much of the internal structure and biochemistry of plastids, for instance the presence of thylakoids and particular chlorophylls, is very similar to that of cyanobacteria. Phylogenetic estimates constructed with bacteria, plastids, and eukaryotic genomes also suggest that plastids are most closely related to cyanobacteria.
Mitochondria have several enzymes and transport systems similar to those of bacteria.
Some proteins encoded in the nucleus are transported to the organelle, and both mitochondria and plastids have small genomes compared to bacteria. This is consistent with an increased dependence on the eukaryotic host after forming an endosymbiosis. Most genes on the organellar genomes have been lost or moved to the nucleus. Most genes needed for mitochondrial and plastid function are located in the nucleus. Many originate from the bacterial endosymbiont.
Plastids are present in very different groups of protists, some of which are closely related to forms lacking plastids. This suggests that if chloroplasts originated de novo, they did so multiple times, in which case their close similarity to each other is difficult to explain.
Many of these protists contain "primary" plastids that have not yet been acquired from other plastid-containing eukaryotes.
Among eukaryotes that acquired their plastids directly from bacteria (known as Archaeplastida), the glaucophyte algae have chloroplasts that strongly resemble cyanobacteria. In particular, they have a peptidoglycan cell wall between the two membranes.

Comparison of chloroplasts and cyanobacteria showing their similarities.

Secondary endosymbiosis

Primary endosymbiosis involves the engulfment of a bacterium by another free living organism. Secondary endosymbiosis occurs when the product of primary endosymbiosis is itself engulfed and retained by another free living eukaryote. Secondary endosymbiosis has occurred several times and has given rise to extremely diverse groups of algae and other eukaryotes. Some organisms can take opportunistic advantage of a similar process, where they engulf an alga and use the products of its photosynthesis, but once the prey item dies (or is lost) the host returns to a free living state. Obligate secondary endosymbionts become dependent on their organelles and are unable to survive in their absence (for a review see McFadden 2001^[31]). RedToL, the Red Algal Tree of Life Initiative funded by the National Science Foundation highlights the role red algae or Rhodophyta played in the evolution of our planet through secondary endosymbiosis.

One possible secondary endosymbiosis in process has been observed by Okamoto & Inouye (2005). The heterotrophic protist Hatena behaves like a predator until it ingests a green alga, which loses its flagella and cytoskeleton, while Hatena, now a host, switches to photosynthetic nutrition, gains the ability to move towards light and loses its feeding apparatus.^[32]

The process of secondary endosymbiosis left its evolutionary signature within the unique topography of plastid membranes. Secondary plastids are surrounded by three (in euglenophytes and some dinoflagellates) or four membranes (in haptophytes, heterokonts, cryptophytes, and chlorarachniophytes). The two additional membranes are thought to correspond to the plasma membrane of the engulfed alga and the phagosomal membrane of the host cell. The endosymbiotic acquisition of a eukaryote cell is represented in the cryptophytes; where the remnant nucleus of the red algal symbiont (the nucleomorph) is present between the two inner and two outer plastid membranes.^{[citation needed]}

Despite the diversity of organisms containing plastids, the morphology, biochemistry, genomic organisation, and molecular phylogeny of plastid RNAs and proteins suggest a single origin of all extant plastids – although this theory is still debated.^[33]^[34]

Some species including Pediculus anus have multiple chromosomes in the mitochondrion. This and the phylogenetics of the genes encoded within the mitochondrion suggest that mitochondria have multiple ancestors, that these were acquired by endosymbiosis on several occasions rather than just once, and that there have been extensive mergers and rearrangements of genes on the several original mitochondrial chromosomes.^[35]

References

^ "Mitochondria Share an Ancestor With SAR11, a Globally Significant Marine Microbe". ScienceDaily. July 25, 2011. Retrieved 2011-07-26.
^ J. Cameron Thrash et al. (2011). "Phylogenomic evidence for a common ancestor of mitochondria and the SAR11 clade". Scientific Reports. doi:10.1038/srep00013.
^ Deusch, O. et al. Genes of cyanobacterial origin in plant nuclear genomes point to a heterocyst-forming plastid ancestor. Mol. Biol. Evol. 25, 748–761 (2008). doi: 10.1093/molbev/msn022
^ Ochoa de Alda JAG, Esteban, R, Diago, ML, Houmard, J. The plastid ancestor originated among one of the major cyanobacterial lineages. Nature Communications, 5 Article number: 4937 doi:10.1038/ncomms5937.
^ Mereschkowsky, Konstantin (1910). "Theorie der zwei Plasmaarten als Grundlage der Symbiogenesis, einer neuen Lehre von der Ent‐stehung der Organismen.". Biol Centralbl. 30: 353‐367.
^ Mereschkowski C (1905). "Über Natur und Ursprung der Chromatophoren im Pflanzenreiche". Biol Centralbl 25: 593–604.
^ Martin, William; Mayo Roettger, Thorsten Kloesges, Thorsten Thiergart, Christian Woehle, Sven Gould and Tal Dagan. "Modern endosymbiotic theory: Getting lateral gene transfer in-to the equation". Journal of Endocytobiosis and Cell Research 23: 1–5. (journal URL: [1])
^ Schimper AFW (1883). "Über die Entwicklung der Chlorophyllkörner und Farbkörper". Bot. Zeitung 41: 105–14, 121–31, 137–46, 153–62.
^ Wallin IE (1923). "The Mitochondria Problem". The American Naturalist 57 (650): 255–61. doi:10.1086/279919.
^ Wallin, I.E. (1927). Symbionticism and the origin of species. Baltimore: Williams & Wilkins Company. p. 171.
^ Margulis, Lynn (2011). "Symbiogenesis. A new principle of evolution rediscovery of Boris Mikhaylovich Kozo-Polyansky (1890–1957)". Paleontological Journal 44 (12): 1525–1539. doi:10.1134/S0031030110120087.
^ Corning, Peter A. (2010). Holistic Darwinism: Synergy, Cybernetics, and the Bioeconomics of Evolution. Chicago: University of Chicago Press. p. 81. ISBN 978-0-22611-633-4.
^ Ris H, Singh RN (January 1961). "Electron microscope studies on blue-green algae". J Biophys Biochem Cytol 9 (1): 63–80. doi:10.1083/jcb.9.1.63. PMC 2224983. PMID 13741827.
^ Stocking C and Gifford E (1959). "Incorporation of thymidine into chloroplasts of Spirogyra". Biochem. Biophys. Res. Comm. 1 (3): 159–64. doi:10.1016/0006-291X(59)90010-5.
^ Lynn Sagan (1967). "On the origin of mitosing cells". J Theor Biol 14 (3): 255–274. doi:10.1016/0022-5193(67)90079-3. PMID 11541392.
^ Margulis, Lynn; Sagan, Dorion (2001). "Marvellous microbes". Resurgence 206: 10–12.
^ Gabaldón T, Snel B, van Zimmeren F, Hemrika W, Tabak H, Huynen MA (2006). "Origin and evolution of the peroxisomal proteome". Biol. Direct 1 (1): 8. doi:10.1186/1745-6150-1-8. PMC 1472686. PMID 16556314. (Provides evidence that contradicts an endosymbiotic origin of peroxisomes. Instead it is suggested that they evolutionarily originate from the Endoplasmic Reticulum)
^ Michael Syvanen, Clarence I. Kado Horizontal Gene Transfer Academic Press, p. 405 ISBN 978-0126801262
^ ^a ^b Keeling, P. J.; Archibald, J.M. (2008). "Organelle evolution: what’s in a name?". Current Biology 18: 345–347. doi:10.1016/j.cub.2008.02.065. PMID 18430636.
^ Nowack, E.C.; Melkonian, M. & Glockner, G. (2008). "Chromatophore genome sequence of Paulinella sheds light on acquisition of photosynthesis by eukaryotes.". Current Biology 18: 410–418. doi:10.1016/j.cub.2008.02.051. PMID 18356055.
^ [2] Kimball, J. 2010. Kimball's Biology Pages. Accessed October 13, 2010. An online open source biology text by Harvard professor, and author of a general biology text, John W. Kimball.
^ Reece, J., Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson, 2010. Campbell Biology. 9th Edition Benjamin Cummings; 9th Ed. (October 7, 2010)
^ Raven, P., George Johnson, Kenneth Mason, Jonathan Losos, Susan Singer, 2010. Biology. McGraw-Hill 9th Ed. (January 14, 2010)
^ Wise, Robert R; Hoober,, J. Kenneth (2007). Structure and function of plastids. Berlin: Springer. p. 104. ISBN 9781402065705.
^ Fischer K, Weber A, Brink S, Arbinger B, Schünemann D, Borchert S, Heldt HW, Popp B, Benz R, Link TA (1994). "Porins from plants. Molecular cloning and functional characterization of two new members of the porin family". J Biol Chem 269 (41): 25754–25760. PMID 7523392.
^ Zeth K, Thein M (2010). "Porins in prokaryotes and eukaryotes: common themes and variations". Biochem J 431 (1): 13–22. doi:10.1042/BJ20100371. PMID 20836765.
^ Fairman JW, Noinaj N, Buchanan SK (2011). "The structural biology of β-barrel membrane proteins: a summary of recent reports". Current Opinion in Structural Biology 21 (4): 523–531. doi:10.1016/j.sbi.2011.05.005. PMC 3164749. PMID 21719274.
^ Mileykovskaya E, Dowhan W (2009). "Cardiolipin membrane domains in prokaryotes and eukaryotes". Biochim Biophys Acta 1788 (10): 2084–2091. doi:10.1016/j.bbamem.2009.04.003. PMC 2757463. PMID 19371718.
^ Andersson SG, Zomorodipour A, Andersson JO, Sicheritz-Pontén T, Alsmark UC, Podowski RM, Näslund AK, Eriksson AS, Winkler HH, Kurland CG (1998). "The genome sequence of Rickettsia prowazekii and the origin of mitochondria". Nature 396 (6707): 133–140. doi:10.1038/24094. PMID 9823893.
^ Dagan T, Roettger M, Stucken K, Landan G, Koch R, Major P, Gould SB, Goremykin VV, Rippka R, Tandeau de Marsac N, Gugger M, Lockhart PJ, Allen JF, Brune I, Maus I, Pühler A, Martin WF (2013). "Genomes of Stigonematalean cyanobacteria (subsection V) and the evolution of oxygenic photosynthesis from prokaryotes to plastids". Genome Biol Evol 5 (1): 31–44. doi:10.1093/gbe/evs117. PMC 3595030. PMID 23221676.
^ McFadden GI (2001). "Primary and secondary endosymbiosis and the origin of plastids". J Phycology 37 (6): 951–9. doi:10.1046/j.1529-8817.2001.01126.x.
^ Okamoto, Noriko; Inouye, Isao (2005). "A Secondary Symbiosis in Progress?". Science 310 (5746): 287. doi:10.1126/science.1116125. Retrieved 15 February 2015.
^ McFadden GI, van Dooren GG (July 2004). "Evolution: red algal genome affirms a common origin of all plastids". Curr. Biol. 14 (13): R514–6. doi:10.1016/j.cub.2004.06.041. PMID 15242632.
^ Gould SB, Waller RF, McFadden GI (2008). "Plastid evolution". Annu Rev Plant Biol 59 (1): 491–517. doi:10.1146/annurev.arplant.59.032607.092915. PMID 18315522.
^ Georgiades, K. and Raoult, D. (2011). "The rhizome of Reclinomonas americana, Homo sapiens, Pediculus humanus and Saccharomyces cerevisiae mitochondria". Biology Direct 6 (55).

External links

Tree of Life Eukaryotes

English Journal

The organization and control of an evolving interdependent population.

Vural DC1, Isakov A2, Mahadevan L3.
Journal of the Royal Society, Interface / the Royal Society.J R Soc Interface.2015 Jul 6;12(108):20150044. doi: 10.1098/rsif.2015.0044.
Starting with Darwin, biologists have asked how populations evolve from a low fitness state that is evolutionarily stable to a high fitness state that is not. Specifically of interest is the emergence of cooperation and multicellularity where the fitness of individuals often appears in conflict with
PMID 26040593

[Symbiogenesis and Synthetic Evolutionary Theory: The Third Synthesis].

Provoro NA, Tikhonovich IA, Vorobyov NI.
Genetika.Genetika.2015 Jun;51(6):658-67.
Integration of the concepts of symbiogenesis and synthetic evolutionary theory is the main path for the development of evolutionary biology. It is based on the analysis of cooperative adaptations that evolve under the impact of symbiotic-specific selective pressures responsible for the formation of
PMID 26310029

[Bacterial Genome Evolution in Superspecies Systems: an Approach to the Reconstruction of Symbiogenesis Processes].

Provorov NA, Tikhonovich IA.
Genetika.Genetika.2015 Apr;51(4):456-65.
Bacteria form a broad spectrum of symbioses with eukaryotes. This permits reconstruction of the symbiogenesis processes providing the transformation of free-living microorganisms into cellular organelles. In ecologically (conditionally) obligate symbioses, an increase in the size and complexity of t
PMID 26087621

Japanese Journal

確率コンセプションの共生発生に関する一考察

数学教育学研究 : 全国数学教育学会誌 17(2), 25-33, 2011
NAID 110009616207

微細藻類の共生進化 (特集黒潮圏の自然と人間の共存・共生(2))

海洋と生物 27(6), 612-617, 2005-12
NAID 40007099326

The ultrastructural features and division of secondary plastids

Journal of plant research 118(3), 163-172, 2005-06-01
NAID 10016578350

Related Pictures

[1] "Mitochondria Share an Ancestor With SAR11, a Globally Significant Marine Microbe". ScienceDaily. July 25, 2011. Retrieved 2011-07-26.

[2] J. Cameron Thrash et al. (2011). "Phylogenomic evidence for a common ancestor of mitochondria and the SAR11 clade". Scientific Reports. doi:10.1038/srep00013.

[3] Deusch, O. et al. Genes of cyanobacterial origin in plant nuclear genomes point to a heterocyst-forming plastid ancestor. Mol. Biol. Evol. 25, 748–761 (2008). doi: 10.1093/molbev/msn022

[4] Ochoa de Alda JAG, Esteban, R, Diago, ML, Houmard, J. The plastid ancestor originated among one of the major cyanobacterial lineages. Nature Communications, 5 Article number: 4937 doi:10.1038/ncomms5937.

[Mereschkowsky2-5] Mereschkowsky, Konstantin (1910). "Theorie der zwei Plasmaarten als Grundlage der Symbiogenesis, einer neuen Lehre von der Ent‐stehung der Organismen.". Biol Centralbl. 30: 353‐367.

[6] Mereschkowski C (1905). "Über Natur und Ursprung der Chromatophoren im Pflanzenreiche". Biol Centralbl 25: 593–604.

[Martin-7] Martin, William; Mayo Roettger, Thorsten Kloesges, Thorsten Thiergart, Christian Woehle, Sven Gould and Tal Dagan. "Modern endosymbiotic theory: Getting lateral gene transfer in-to the equation". Journal of Endocytobiosis and Cell Research 23: 1–5. (journal URL: [1])

[8] Schimper AFW (1883). "Über die Entwicklung der Chlorophyllkörner und Farbkörper". Bot. Zeitung 41: 105–14, 121–31, 137–46, 153–62.

[9] Wallin IE (1923). "The Mitochondria Problem". The American Naturalist 57 (650): 255–61. doi:10.1086/279919.

[WallinBk-10] Wallin, I.E. (1927). Symbionticism and the origin of species. Baltimore: Williams & Wilkins Company. p. 171.

[11] Margulis, Lynn (2011). "Symbiogenesis. A new principle of evolution rediscovery of Boris Mikhaylovich Kozo-Polyansky (1890–1957)". Paleontological Journal 44 (12): 1525–1539. doi:10.1134/S0031030110120087.

[12] Corning, Peter A. (2010). Holistic Darwinism: Synergy, Cybernetics, and the Bioeconomics of Evolution. Chicago: University of Chicago Press. p. 81. ISBN 978-0-22611-633-4.

[13] Ris H, Singh RN (January 1961). "Electron microscope studies on blue-green algae". J Biophys Biochem Cytol 9 (1): 63–80. doi:10.1083/jcb.9.1.63. PMC 2224983. PMID 13741827.

[14] Stocking C and Gifford E (1959). "Incorporation of thymidine into chloroplasts of Spirogyra". Biochem. Biophys. Res. Comm. 1 (3): 159–64. doi:10.1016/0006-291X(59)90010-5.

[15] Lynn Sagan (1967). "On the origin of mitosing cells". J Theor Biol 14 (3): 255–274. doi:10.1016/0022-5193(67)90079-3. PMID 11541392.

[16] Margulis, Lynn; Sagan, Dorion (2001). "Marvellous microbes". Resurgence 206: 10–12.

[17] Gabaldón T, Snel B, van Zimmeren F, Hemrika W, Tabak H, Huynen MA (2006). "Origin and evolution of the peroxisomal proteome". Biol. Direct 1 (1): 8. doi:10.1186/1745-6150-1-8. PMC 1472686. PMID 16556314. (Provides evidence that contradicts an endosymbiotic origin of peroxisomes. Instead it is suggested that they evolutionarily originate from the Endoplasmic Reticulum)

[18] Michael Syvanen, Clarence I. Kado Horizontal Gene Transfer Academic Press, p. 405 ISBN 978-0126801262

[Keeling_2008_345.E2.80.93347-19] Keeling, P. J.; Archibald, J.M. (2008). "Organelle evolution: what’s in a name?". Current Biology 18: 345–347. doi:10.1016/j.cub.2008.02.065. PMID 18430636.

[20] Nowack, E.C.; Melkonian, M. & Glockner, G. (2008). "Chromatophore genome sequence of Paulinella sheds light on acquisition of photosynthesis by eukaryotes.". Current Biology 18: 410–418. doi:10.1016/j.cub.2008.02.051. PMID 18356055.

[Kimball-21] [2] Kimball, J. 2010. Kimball's Biology Pages. Accessed October 13, 2010. An online open source biology text by Harvard professor, and author of a general biology text, John W. Kimball.

[Reece-22] Reece, J., Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson, 2010. Campbell Biology. 9th Edition Benjamin Cummings; 9th Ed. (October 7, 2010)

[Raven-23] Raven, P., George Johnson, Kenneth Mason, Jonathan Losos, Susan Singer, 2010. Biology. McGraw-Hill 9th Ed. (January 14, 2010)

[24] Wise, Robert R; Hoober,, J. Kenneth (2007). Structure and function of plastids. Berlin: Springer. p. 104. ISBN 9781402065705.

[25] Fischer K, Weber A, Brink S, Arbinger B, Schünemann D, Borchert S, Heldt HW, Popp B, Benz R, Link TA (1994). "Porins from plants. Molecular cloning and functional characterization of two new members of the porin family". J Biol Chem 269 (41): 25754–25760. PMID 7523392.

[26] Zeth K, Thein M (2010). "Porins in prokaryotes and eukaryotes: common themes and variations". Biochem J 431 (1): 13–22. doi:10.1042/BJ20100371. PMID 20836765.

[27] Fairman JW, Noinaj N, Buchanan SK (2011). "The structural biology of β-barrel membrane proteins: a summary of recent reports". Current Opinion in Structural Biology 21 (4): 523–531. doi:10.1016/j.sbi.2011.05.005. PMC 3164749. PMID 21719274.

[28] Mileykovskaya E, Dowhan W (2009). "Cardiolipin membrane domains in prokaryotes and eukaryotes". Biochim Biophys Acta 1788 (10): 2084–2091. doi:10.1016/j.bbamem.2009.04.003. PMC 2757463. PMID 19371718.

[29] Andersson SG, Zomorodipour A, Andersson JO, Sicheritz-Pontén T, Alsmark UC, Podowski RM, Näslund AK, Eriksson AS, Winkler HH, Kurland CG (1998). "The genome sequence of Rickettsia prowazekii and the origin of mitochondria". Nature 396 (6707): 133–140. doi:10.1038/24094. PMID 9823893.

[30] Dagan T, Roettger M, Stucken K, Landan G, Koch R, Major P, Gould SB, Goremykin VV, Rippka R, Tandeau de Marsac N, Gugger M, Lockhart PJ, Allen JF, Brune I, Maus I, Pühler A, Martin WF (2013). "Genomes of Stigonematalean cyanobacteria (subsection V) and the evolution of oxygenic photosynthesis from prokaryotes to plastids". Genome Biol Evol 5 (1): 31–44. doi:10.1093/gbe/evs117. PMC 3595030. PMID 23221676.

[31] McFadden GI (2001). "Primary and secondary endosymbiosis and the origin of plastids". J Phycology 37 (6): 951–9. doi:10.1046/j.1529-8817.2001.01126.x.

[32] Okamoto, Noriko; Inouye, Isao (2005). "A Secondary Symbiosis in Progress?". Science 310 (5746): 287. doi:10.1126/science.1116125. Retrieved 15 February 2015.

[33] McFadden GI, van Dooren GG (July 2004). "Evolution: red algal genome affirms a common origin of all plastids". Curr. Biol. 14 (13): R514–6. doi:10.1016/j.cub.2004.06.041. PMID 15242632.

[34] Gould SB, Waller RF, McFadden GI (2008). "Plastid evolution". Annu Rev Plant Biol 59 (1): 491–517. doi:10.1146/annurev.arplant.59.032607.092915. PMID 18315522.

[35] Georgiades, K. and Raoult, D. (2011). "The rhizome of Reclinomonas americana, Homo sapiens, Pediculus humanus and Saccharomyces cerevisiae mitochondria". Biology Direct 6 (55).

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