Archaeplastida | |
---|---|
![]() | |
Trees, grasses and algae in and around Sprague River, Oregon | |
Scientific classification ![]() | |
Domain: | Eukaryota |
(unranked): | Diaphoretickes |
(unranked): | Archaeplastida Adl et al., 2005[1] |
Subgroups | |
Synonyms | |
|
The Archaeplastida (or kingdom Plantae sensu lato "in a broad sense"; pronounced /ɑːrkɪ'plastɪdə/) are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor group glaucophytes.[6] It also includes the non-photosynthetic lineage Rhodelphidia, a predatorial (eukaryotrophic) flagellate that is sister to the Rhodophyta, and probably the microscopic picozoans.[7] The Archaeplastida have chloroplasts that are surrounded by two membranes, suggesting that they were acquired directly through a single endosymbiosis event by feeding on a cyanobacterium.[8] All other groups which have chloroplasts, besides the amoeboid genus Paulinella, have chloroplasts surrounded by three or four membranes, suggesting they were acquired secondarily from red or green algae.[note 1] Unlike red and green algae, glaucophytes have never been involved in secondary endosymbiosis events.[10]
The cells of the Archaeplastida typically lack centrioles and have mitochondria with flat cristae. They usually have a cell wall that contains cellulose, and food is stored in the form of starch. However, these characteristics are also shared with other eukaryotes. The main evidence that the Archaeplastida form a monophyletic group comes from genetic studies, which indicate their plastids probably had a single origin. This evidence is disputed.[11][12] Based on the evidence to date, it is not possible to confirm or refute alternative evolutionary scenarios to a single primary endosymbiosis.[13] Photosynthetic organisms with plastids of different origin (such as brown algae) do not belong to the Archaeplastida.
The archaeplastidans fall into two main evolutionary lines. The red algae are pigmented with chlorophyll a and phycobiliproteins, like most cyanobacteria, and accumulate starch outside the chloroplasts. The green algae and land plants – together known as Viridiplantae (Latin for "green plants") or Chloroplastida – are pigmented with chlorophylls a and b, but lack phycobiliproteins, and starch is accumulated inside the chloroplasts.[14] The glaucophytes have typical cyanobacterial pigments, and are unusual in retaining a cell wall within their plastids (called cyanelles).[1]
Archaeplastida should not be confused with the older and obsolete name Archiplastideae, which refers to cyanobacteria and other groups of bacteria.[15][16]
Taxonomy
The consensus in 2005, when the group consisting of the glaucophytes and red and green algae and land plants was named 'Archaeplastida',[1] was that it was a clade, i.e. was monophyletic. Many studies published since then have provided evidence in agreement.[17][18][19][20] Other studies, though, have suggested that the group is paraphyletic.[21][22][23][12] To date, the situation appears unresolved, but a strong signal for Plantae (Archaeplastida) monophyly has been demonstrated in a recent study (with an enrichment of red algal genes).[24] The assumption made here is that Archaeplastida is a valid clade.
Various names have been given to the group. Some authors have simply referred to the group as plants or Plantae.[25][26] However, the name Plantae is ambiguous, since it has also been applied to less inclusive clades, such as Viridiplantae and embryophytes. To distinguish, the larger group is sometimes known as Plantae sensu lato ("plants in the broad sense").
To avoid ambiguity, other names have been proposed. Primoplantae, which appeared in 2004, seems to be the first new name suggested for this group.[5] Another name applied to this node is Plastida, defined as the clade sharing "plastids of primary (direct prokaryote) origin [as] in Magnolia virginiana Linnaeus 1753".[27]
Although many studies have suggested the Archaeplastida form a monophyletic group,[28] a 2009 paper argues that they are in fact paraphyletic.[23] The enrichment of novel red algal genes in a recent study demonstrates a strong signal for Plantae (Archaeplastida) monophyly and an equally strong signal of gene sharing history between the red/green algae and other lineages.[24] This study provides insight on how rich mesophilic red algal gene data are crucial for testing controversial issues in eukaryote evolution and for understanding the complex patterns of gene inheritance in protists.
The name Archaeplastida was proposed in 2005 by a large international group of authors (Adl et al.), who aimed to produce a classification for the eukaryotes which took into account morphology, biochemistry, and phylogenetics, and which had "some stability in the near term." They rejected the use of formal taxonomic ranks in favour of a hierarchical arrangement where the clade names do not signify rank. Thus, the phylum name 'Glaucophyta' and the class name 'Rhodophyceae' appear at the same level in their classification. The divisions proposed for the Archaeplastida are shown below in both tabular and diagrammatic form.[1]
Archaeplastida:
- Glaucophyta Skuja, 1954 (Glaucocystophyta Kies & Kremer, 1986) – glaucophytes
- Glaucophytes are a small group of freshwater single-celled algae. Their chloroplasts, called cyanelles, have a peptidoglycan layer, making them more similar to cyanobacteria than those of the remaining Archaeplastida.
- Rhodophyceae Thuret, 1855, emend. Rabenhorst, 1863, emend. Adl et al., 2005 (Rhodophyta Wettstein 1901) – red algae
- Red algae form one of the largest groups of algae. Most are seaweeds, being multicellular and marine. Their red colour comes from phycobiliproteins, used as accessory pigments in light capture for photosynthesis.
- Chloroplastida Adl et al., 2005 (Viridiplantae Cavalier-Smith 1981; Chlorobionta Jeffrey 1982, emend. Bremer 1985, emend. Lewis and McCourt 2004; Chlorobiota Kendrick and Crane 1997)
- Chloroplastida is the term chosen by Adl et al. for the group made up of the green algae and land plants (embryophytes). Except where lost secondarily, all have chloroplasts without a peptidoglycan layer and lack phycobiliproteins.
- Chlorophyta Pascher, 1914, emend. Lewis & McCourt, 2004 – green algae (part)
- Adl et al. employ a narrow definition of the Chlorophyta; other sources include the Chlorodendrales and Prasinophytae, which may themselves be combined.
- Ulvophyceae Mattox & Stewart, 1984
- Trebouxiophyceae Friedl, 1995 (Pleurastrophyceae Mattox et al. 1984; Microthamniales Melkonian 1990)
- Chlorophyceae Christensen, 1994
- Chlorodendrales Fritsch, 1917 – green algae (part)
- Prasinophytae Cavalier-Smith, 1998, emend. Lewis & McCourt, 2004 – green algae (part)
- Mesostigma Lauterborn, 1894, emend. McCourt in Adl et al., 2005 (Mesostigmata Turmel, Otis, and Lemieux 2002)
- Charophyta Karol et al., 2001, emend. Lewis & McCourt, 2004 (Charophyceae Smith 1938, emend. Mattox and Stewart 1984) – green algae (part) and land plants
- Charophyta sensu lato, as used by Adl et al., is a monophyletic group which is made up of some green algae, including the stoneworts (Charophyta sensu stricto), as well as the land plants (embryophytes).
- Sub-divisions other than Streptophytina (below) were not given by Adl et al.
- Other sources would include the green algal groups Chlorokybales, Klebsormidiales, Zygnematales and Coleochaetales.[29]
- Streptophytina Lewis & McCourt, 2004 – stoneworts and land plants
Cladogram
Below is a consensus reconstruction of green algal relationships, mainly based on molecular data.[30][31][32][33][34][35][36][37][38][39] While the Glaucophyta are typically figured as deepest rooting Archeaplastida,[40][41][42][43] some genomic research points to Rhodophyta as basal, possibly with Cryptista and picozoa emerging in Archaeplastida.[44][45][46][47][48][49][50][51][3] At least for Cryptista, this analysis (by Burki et al 2016) has gained credibility.[52][53][54][55][56]
|
However, there is a lot of contention near the Archaeplastida root, e.g. whether Glaucophyta or Rhodophyta are basal, or whether e.g. Cryptista emerged within the Archaeplastida. In 2014 a thorough review was published on these inconsistencies.[58] The position of Telonemia and Picozoa are not clear. Also Hacrobia (Haptista + Cryptista) may be completely associated with the SAR clade. The SAR are often seen as eukaryote-eukaryote hybrids, contributing to the confusion in the genetic analyses. A sister of Gloeomargarita lithophora has been engulfed by an ancestor of the Archaeplastida, leading to the plastids which are living in permanent endosymbiosis in most of the descendent lineages. Because both Gloeomargarita and related cyanobacteria, in addition to the most primitive archaeplastids, all live in freshwater, it seems the Archaeplastida originated in freshwater, and only colonized the oceans in the late Proterozoic.[59][60]
Morphology
All archaeplastidans have plastids (chloroplasts) that carry out photosynthesis and are believed to be derived from endosymbiotic cyanobacteria. In glaucophytes, perhaps the most primitive members of the group, the chloroplast is called a cyanelle and shares several features with cyanobacteria, including a peptidoglycan cell wall, that are not retained in other members of the group. The resemblance of cyanelles to cyanobacteria supports the endosymbiotic theory.
The cells of most archaeplastidans have walls, commonly but not always made of cellulose.
The Archaeplastida vary widely in the degree of their cell organization, from isolated cells to filaments to colonies to multi-celled organisms. The earliest were unicellular, and many groups remain so today. Multicellularity evolved separately in several groups, including red algae, ulvophyte green algae, and in the green algae that gave rise to stoneworts and land plants.
Endosymbiosis
Because the ancestral archaeplastidan is hypothesized to have acquired its chloroplasts directly by engulfing cyanobacteria, the event is known as a primary endosymbiosis (as reflected in the name chosen for the group 'Archaeplastida' i.e. 'ancient plastid'). One species of green algae, Cymbomonas tetramitiformis in the order Pyramimonadales, is a mixotroph and able to support itself through both phagotrophy and phototrophy. It is not yet known if this is a primitive trait and therefore defines the last common ancestor of Archaeplastida, which could explain how it obtained its chloroplasts, or if it is a trait regained by horizontal gene transfer.[61]
Evidence for primary endosymbosis includes the presence of a double membrane around the chloroplasts; one membrane belonged to the bacterium, and the other to the eukaryote that captured it. Over time, many genes from the chloroplast have been transferred to the nucleus of the host cell. The presence of such genes in the nuclei of eukaryotes without chloroplasts suggests this transfer happened early in the evolution of the group.[62]
Other eukaryotes with chloroplasts appear to have gained them by engulfing a single-celled archaeplastidan with its own bacterially-derived chloroplasts. Because these events involve endosymbiosis of cells that have their own endosymbionts, the process is called secondary endosymbiosis. The chloroplasts of such eukaryotes are typically surrounded by more than two membranes, reflecting a history of multiple engulfment. The chloroplasts of euglenids, chlorarachniophytes and a small group of dinoflagellates appear to be captured green algae,[63] whereas those of the remaining photosynthetic eukaryotes, such as heterokont algae, cryptophytes, haptophytes, and dinoflagellates, appear to be captured red algae.
Fossil record
Perhaps the most ancient remains of Archaeplastida are putative red algae (Rafatazmia) within stromatolites in 1600 Ma (million years ago) rocks in India.[64] Somewhat more recent are microfossils from the Roper group in northern Australia. The structure of these single-celled fossils resembles that of modern green algae. They date to the Mesoproterozoic Era, about 1500 to 1300 Ma.[65] These fossils are consistent with a molecular clock study that calculated that this clade diverged about 1500 Ma.[66] The oldest fossil that can be assigned to a specific modern group is the red alga Bangiomorpha, from 1200 Ma.[67]
In the late Neoproterozoic Era, algal fossils became more numerous and diverse. Eventually, in the Paleozoic Era, plants emerged onto land, and have continued to flourish up to the present.
Notes
- ^ The exceptional two plastid membranes of the stramenopile alga Chrysoparadoxa are probably the result of secondary reduction.[9]
References
- ^ a b c d Adl, S.M.; et al. (2005). "The New Higher Level Classification of Eukaryotes with Emphasis on the Taxonomy of Protists". Journal of Eukaryotic Microbiology. 52 (5): 399–451. doi:10.1111/j.1550-7408.2005.00053.x. PMID 16248873. S2CID 8060916.
- ^ Yazaki, Euki; Yabuki, Akinori; Imaizumi, Ayaka; Kume, Keitaro; Hashimoto, Tetsuo; Inagaki, Yuji (31 August 2021). "Phylogenomics invokes the clade housing Cryptista, Archaeplastida, and Microheliella maris". doi:10.1101/2021.08.29.458128. S2CID 237393558. Retrieved 25 November 2021.
{{cite journal}}
: Cite journal requires|journal=
(help) - ^ a b Schön, M.E.; Zlatogursky, V.V.; Singh, R.P.; et al. (17 November 2021). "Single cell genomics reveals plastid-lacking Picozoa are close relatives of red algae". Nature Communications. 12: 6651 (1): 6651. Bibcode:2021NatCo..12.6651S. doi:10.1038/s41467-021-26918-0. ISSN 2041-1723. PMC 8599508. PMID 34789758.
- ^ Cavalier-Smith, T. (1981). "Eukaryote Kingdoms: Seven or Nine?"". BioSystems. 14 (3–4): 461–481. doi:10.1016/0303-2647(81)90050-2. PMID 7337818.
- ^ a b Palmer, Jeffrey D.; Soltis, Douglas E.; Chase, Mark W. (2004). "The plant tree of life: an overview and some points of view". American Journal of Botany. 91 (10): 1437–1445. doi:10.3732/ajb.91.10.1437. PMID 21652302.
- ^ Ball, S.; Colleoni, C. (January 2011). Cenci, U.; Raj, J.N.; Tirtiaux, C. "The evolution of glycogen and starch metabolism in eukaryotes gives molecular clues to understand the establishment of plastid endosymbiosis". Journal of Experimental Botany. 62 (6): 1775–1801. doi:10.1093/jxb/erq411. PMID 21220783.
- ^ Picozoans Are Algae After All: Study | The Scientist Magazine®
- ^ Papanin Institute for Biology of Inland Waters, Russian Academy of Sciences; Tikhonenkov, Denis V. (2020). "Predatory flagellates – the new recently discovered deep branches of the eukaryotic tree and their evolutionary and ecological significance" (PDF). Protistology. 14 (1). doi:10.21685/1680-0826-2020-14-1-2.
- ^ Wetherbee, Richard; Jackson, Christopher J.; Repetti, Sonja I.; Clementson, Lesley A.; Costa, Joana F.; van de Meene, Allison; Crawford, Simon; Verbruggen, Heroen (9 December 2018). "The golden paradox – a new heterokont lineage with chloroplasts surrounded by two membranes". Journal of Phycology. 22 (2): 257–278. doi:10.1111/jpy.12822. hdl:11343/233613. PMID 30536815. S2CID 54477112.
- ^ Handbook of Marine Microalgae: Biotechnology Advances
- ^ Parfrey. L. W.; Barbero, E.; Lasser, E; et al. (December 2006). "Evaluating support for the current classification of eukaryotic diversity". PLOS Genetics. 2 (12): e220. doi:10.1371/journal.pgen.0020220. PMC 1713255. PMID 17194223.
- ^ a b Kim, E; Graham, L. E. (July 2008). Redfield, Rosemary Jeanne (ed.). "EEF2 analysis challenges the monophyly of Archaeplastida and Chromalveolata". PLOS ONE. 3 (7): e2621. Bibcode:2008PLoSO...3.2621K. doi:10.1371/journal.pone.0002621. PMC 2440802. PMID 18612431.
- ^ Mackiewicz, P.; Gagat, P. (2014). "Monophyly of Archaeplastida supergroup and relationships among its lineages in the light of phylogenetic and phylogenomic studies. Are we close to a consensus?". Acta Societatis Botanicorum Poloniae. 83 (4): 263–280. doi:10.5586/asbp.2014.044.
- ^ Viola, R.; Nyvall, P.; Pedersén, M. (2001). "The unique features of starch metabolism in red algae". Proceedings of the Royal Society B: Biological Sciences. 268 (1474): 1417–1422. doi:10.1098/rspb.2001.1644. PMC 1088757. PMID 11429143.
- ^ Copeland, H. F. (1956). The Classification of Lower Organisms. Palo Alto: Pacific Books, p. 29, [1].
- ^ Bessey, C. E. (1907). "A Synopsis of Plant Phyla". Univ. Nebraska Studies. 7: 275–358.
- ^ Burki, Fabien; Kamran Shalchian-Tabrizi; Marianne Minge; Åsmund Skjæveland; Sergey I. Nikolaev; Kjetill S. Jakobsen; Jan Pawlowski (2007). Butler, Geraldine (ed.). "Phylogenomics Reshuffles the Eukaryotic Supergroups". PLOS ONE. 2 (8): e790. Bibcode:2007PLoSO...2..790B. doi:10.1371/journal.pone.0000790. PMC 1949142. PMID 17726520.
- ^ Burki, F.; Inagaki, Y.; Brate, J.; Archibald, J. M.; Keeling, P. J.; Cavalier-Smith, T.; et al. (2009). "Large-Scale Phylogenomic Analyses Reveal That Two Enigmatic Protist Lineages, Telonemia and Centroheliozoa, Are Related to Photosynthetic Chromalveolates". Genome Biology and Evolution. 1: 231–238. doi:10.1093/gbe/evp022. PMC 2817417. PMID 20333193.
- ^ Cavalier-Smith, Thomas (2009). "Kingdoms Protozoa and Chromista and the eozoan root of the eukaryotic tree". Biology Letters. 6 (3): 342–345. doi:10.1098/rsbl.2009.0948. PMC 2880060. PMID 20031978.
- ^ Rogozin, I. B.; Basu, M. K.; Csürös, M. & Koonin, E. V. (2009). "Analysis of Rare Genomic Changes Does Not Support the Unikont–Bikont Phylogeny and Suggests Cyanobacterial Symbiosis as the Point of Primary Radiation of Eukaryotes". Genome Biology and Evolution. 1: 99–113. doi:10.1093/gbe/evp011. PMC 2817406. PMID 20333181.
- ^ Baldauf, S.L., Roger, A.J., Wenk-Siefert, I., Doolittle, W.F. 2000. A kingdom-level phylogeny of eukaryotes based on combined protein data. Science 290: 972-977 (researchgate.net)
- ^ Lipscomb, Diana. 1991. Broad classification: the kingdoms and th protozoa. In: Parasitic Protozoa, Vol. 1, 2nd ed., J.P. Kreier, J.R. Baker (eds.), pp. 81-136. Academic Press, San Diego.
- ^ a b Nozaki, H.; Maruyama, S.; Matsuzaki, M.; Nakada, T.; Kato, S.; Misawa, K. (December 2009). "Phylogenetic positions of Glaucophyta, green plants (Archaeplastida) and Haptophyta (Chromalveolata) as deduced from slowly evolving nuclear genes". Molecular Phylogenetics and Evolution. 53 (3): 872–80. doi:10.1016/j.ympev.2009.08.015. PMID 19698794.
- ^ a b Chan, C. X.; Yang, E. C.; Banerjee, T.; Yoon, H. S.; Martone, P. T.; Estevez, J. M.; Bhattacharya, D. (2011). "Red and green algal monophyly and extensive gene sharing found in a rich repertoire of red algal genes". Current Biology. 21 (4): 328–333. doi:10.1016/j.cub.2011.01.037. PMID 21315598. S2CID 7162977.
- ^ T. Cavalier-Smith (1981). "Eukaryote Kingdoms: Seven or Nine?". BioSystems. 14 (3–4): 461–481. doi:10.1016/0303-2647(81)90050-2. PMID 7337818.
- ^ Bhattacharya, Debashish; Yoon, Hwan Su; Hackett, Jeremiah (2003). "Photosynthetic eukaryotes unite: endosymbiosis connects the dots". BioEssays. 26 (1): 50–60. doi:10.1002/bies.10376. PMID 14696040.
- ^ Simpson, A. G. B. (2004). "Highest-level taxa within Eukaryotes". First International Phylogenetic Nomenclature Meeting. Paris, July 6–9.
- ^ Vinogradov S. N.; Fernández, I.; Hoogewijs, D.; Arredondo-Peter, R. (October 2010). "Phylogenetic Relationships of 3/3 and 2/2 Hemoglobins in Archaeplastida Genomes to Bacterial and Other Eukaryote Hemoglobins". Molecular Plant. 4 (1): 42–58. doi:10.1093/mp/ssq040. PMID 20952597.
- ^ Turmel, M.; Otis, C.; Lemieux, C. (2005). "The complete chloroplast DNA sequences of the charophycean green algae Staurastrum and Zygnema reveal that the chloroplast genome underwent extensive changes during the evolution of the Zygnematales". BMC Biology. 3: 22. doi:10.1186/1741-7007-3-22. PMC 1277820. PMID 16236178.
- ^ Leliaert, Frederik; Smith, David R.; Moreau, Hervé; Herron, Matthew D.; Verbruggen, Heroen; Delwiche, Charles F.; De Clerck, Olivier (2012). "Phylogeny and Molecular Evolution of the Green Algae" (PDF). Critical Reviews in Plant Sciences. 31: 1–46. doi:10.1080/07352689.2011.615705. S2CID 17603352.
- ^ Marin, Birger (2012). "Nested in the Chlorellales or Independent Class? Phylogeny and Classification of the Pedinophyceae (Viridiplantae) Revealed by Molecular Phylogenetic Analyses of Complete Nuclear and Plastid-encoded rRNA Operons". Protist. 163 (5): 778–805. doi:10.1016/j.protis.2011.11.004. PMID 22192529.
- ^ Laurin-Lemay, Simon; Brinkmann, Henner; Philippe, Hervé (2012). "Origin of land plants revisited in the light of sequence contamination and missing data". Current Biology. 22 (15): R593–R594. doi:10.1016/j.cub.2012.06.013. PMID 22877776.
- ^ Leliaert, Frederik; Tronholm, Ana; Lemieux, Claude; Turmel, Monique; DePriest, Michael S.; Bhattacharya, Debashish; Karol, Kenneth G.; Fredericq, Suzanne; Zechman, Frederick W. (2016-05-09). "Chloroplast phylogenomic analyses reveal the deepest-branching lineage of the Chlorophyta, Palmophyllophyceae class. nov". Scientific Reports. 6: 25367. Bibcode:2016NatSR...625367L. doi:10.1038/srep25367. ISSN 2045-2322. PMC 4860620. PMID 27157793.
- ^ Cook, Martha E.; Graham, Linda E. (2017). Archibald, John M.; Simpson, Alastair G. B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Springer International Publishing. pp. 185–204. doi:10.1007/978-3-319-28149-0_36. ISBN 9783319281476.
- ^ Lewis, Louise A.; Richard M. McCourt (2004). "Green algae and the origin of land plants". American Journal of Botany. 91 (10): 1535–1556. doi:10.3732/ajb.91.10.1535. PMID 21652308. Archived from the original (abstract) on 2011-02-21. Retrieved 2017-10-15.
- ^ Ruhfel, Brad R.; Gitzendanner, Matthew A.; Soltis, Pamela S.; Soltis, Douglas E.; Burleigh, J. Gordon (2014-02-17). "From algae to angiosperms–inferring the phylogeny of green plants (Viridiplantae) from 360 plastid genomes". BMC Evolutionary Biology. 14: 23. doi:10.1186/1471-2148-14-23. ISSN 1471-2148. PMC 3933183. PMID 24533922.
- ^ Adl, Sina M.; Simpson, Alastair G. B.; Lane, Christopher E.; Lukeš, Julius; Bass, David; Bowser, Samuel S.; Brown, Matthew W.; Burki, Fabien; Dunthorn, Micah (2012-09-01). "The Revised Classification of Eukaryotes". Journal of Eukaryotic Microbiology. 59 (5): 429–514. doi:10.1111/j.1550-7408.2012.00644.x. ISSN 1550-7408. PMC 3483872. PMID 23020233.
- ^ Lemieux, Claude; Otis, Christian; Turmel, Monique (2007-01-12). "A clade uniting the green algae Mesostigma viride and Chlorokybus atmophyticus represents the deepest branch of the Streptophyta in chloroplast genome-based phylogenies". BMC Biology. 5: 2. doi:10.1186/1741-7007-5-2. ISSN 1741-7007. PMC 1781420. PMID 17222354.
- ^ Umen, James G. (2014-11-01). "Green Algae and the Origins of Multicellularity in the Plant Kingdom". Cold Spring Harbor Perspectives in Biology. 6 (11): a016170. doi:10.1101/cshperspect.a016170. ISSN 1943-0264. PMC 4413236. PMID 25324214.
- ^ Vries, Jan de; Gould, Sven B. (2017-01-01). "The monoplastidic bottleneck in algae and plant evolution". J Cell Sci. 131 (2): jcs.203414. doi:10.1242/jcs.203414. ISSN 0021-9533. PMID 28893840.
- ^ Ponce-Toledo, Rafael I.; Deschamps, Philippe; López-García, Purificación; Zivanovic, Yvan; Benzerara, Karim; Moreira, David (2017). "An Early-Branching Freshwater Cyanobacterium at the Origin of Plastids". Current Biology. 27 (3): 386–391. doi:10.1016/j.cub.2016.11.056. PMC 5650054. PMID 28132810.
- ^ Dittami, Simon M.; Heesch, Svenja; Olsen, Jeanine L.; Collén, Jonas (2017-08-01). "Transitions between marine and freshwater environments provide new clues about the origins of multicellular plants and algae" (PDF). Journal of Phycology. 53 (4): 731–745. doi:10.1111/jpy.12547. ISSN 1529-8817. PMID 28509401. S2CID 43325392.
- ^ Smith, David R. (2018). "Lost in the Light: Plastid Genome Evolution in Nonphotosynthetic Algae". Plastid Genome Evolution. Advances in Botanical Research. Vol. 85. pp. 29–53. doi:10.1016/bs.abr.2017.10.001. ISBN 9780128134573.
- ^ Burki, Fabien; Kaplan, Maia; Tikhonenkov, Denis V.; Zlatogursky, Vasily; Minh, Bui Quang; Radaykina, Liudmila V.; Smirnov, Alexey; Mylnikov, Alexander P.; Keeling, Patrick J. (2016-01-27). "Untangling the early diversification of eukaryotes: a phylogenomic study of the evolutionary origins of Centrohelida, Haptophyta and Cryptista". Proc. R. Soc. B. 283 (1823): 20152802. doi:10.1098/rspb.2015.2802. ISSN 0962-8452. PMC 4795036. PMID 26817772.
- ^ Brown, Matthew W.; Heiss, Aaron; Kamikawa, Ryoma; Inagaki, Yuji; Yabuki, Akinori; Tice, Alexander K.; Shiratori, Takashi; Ishida, Ken; Hashimoto, Tetsuo (2017-12-03). "Phylogenomics places orphan protistan lineages in a novel eukaryotic super-group". Genome Biol Evol. 10 (2): 427–433. doi:10.1093/gbe/evy014. PMC 5793813. PMID 29360967.
- ^ Lee, JunMo; Cho, Chung Hyun; Park, Seung In; Choi, Ji Won; Song, Hyun Suk; West, John A.; Bhattacharya, Debashish; Yoon, Hwan Su (2016-09-02). "Parallel evolution of highly conserved plastid genome architecture in red seaweeds and seed plants". BMC Biology. 14: 75. doi:10.1186/s12915-016-0299-5. ISSN 1741-7007. PMC 5010701. PMID 27589960.
- ^ Bodył, Andrzej (2018). "Did some red alga-derived plastids evolve via kleptoplastidy? A hypothesis". Biological Reviews. 93 (1): 201–222. doi:10.1111/brv.12340. ISSN 1464-7931. PMID 28544184. S2CID 24613863.
- ^ Gitzendanner, Matthew A.; Soltis, Pamela S.; Wong, Gane K.-S.; Ruhfel, Brad R.; Soltis, Douglas E. (2018). "Plastid phylogenomic analysis of green plants: A billion years of evolutionary history". American Journal of Botany. 105 (3): 291–301. doi:10.1002/ajb2.1048. ISSN 0002-9122. PMID 29603143.
- ^ Strassert, Jürgen F H; Jamy, Mahwash; Mylnikov, Alexander P; Tikhonenkov, Denis V; Burki, Fabien (2019-01-22). Shapiro, Beth (ed.). "New Phylogenomic Analysis of the Enigmatic Phylum Telonemia Further Resolves the Eukaryote Tree of Life". Molecular Biology and Evolution. Oxford University Press (OUP). 36 (4): 757–765. bioRxiv 10.1101/403329. doi:10.1093/molbev/msz012. ISSN 0737-4038. PMC 6844682. PMID 30668767.
- ^ Gawryluk, Ryan M. R.; Tikhonenkov, Denis V.; Hehenberger, Elisabeth; Husnik, Filip; Mylnikov, Alexander P.; Keeling, Patrick J. (2019-09-08). "Non-photosynthetic predators are sister to red algae". Nature. 572 (7768): 240–243. doi:10.1038/s41586-019-1398-6. ISSN 0028-0836. PMID 31316212. S2CID 197542583.
- ^ Irisarri, Iker; Strassert, Jürgen F H; Burki, Fabien (2021-12-16). Folk, Ryan (ed.). "Phylogenomic Insights into the Origin of Primary Plastids". Systematic Biology. 71 (1): 105–120. doi:10.1093/sysbio/syab036. ISSN 1063-5157. PMID 33988690.
- ^ Blaby-Haas, Crysten E.; Merchant, Sabeeha S. (2019-04-29). "Comparative and Functional Algal Genomics". Annual Review of Plant Biology. Annual Reviews. 70 (1): 605–638. doi:10.1146/annurev-arplant-050718-095841. ISSN 1543-5008. OSTI 1502797. PMID 30822111. S2CID 73512522.
- ^ Nowack, Eva C.M.; Weber, Andreas P.M. (2018-04-29). "Genomics-Informed Insights into Endosymbiotic Organelle Evolution in Photosynthetic Eukaryotes". Annual Review of Plant Biology. Annual Reviews. 69 (1): 51–84. doi:10.1146/annurev-arplant-042817-040209. ISSN 1543-5008. PMID 29489396.
- ^ Strassert, Jürgen F. H.; Irisarri, Iker; Williams, Tom A.; Burki, Fabien (2021-03-25). "A molecular timescale for eukaryote evolution with implications for the origin of red algal-derived plastids". Nature Communications. 12 (1): 1879. Bibcode:2021NatCo..12.1879S. doi:10.1038/s41467-021-22044-z. ISSN 2041-1723. PMC 7994803. PMID 33767194.
- ^ Padilla-Mejia, Norma E.; Makarov, Alexandr A.; Barlow, Lael D.; Butterfield, Erin R.; Field, Mark C. (2021-01-01). "Evolution and diversification of the nuclear envelope". Nucleus. 12 (1): 21–41. doi:10.1080/19491034.2021.1874135. ISSN 1949-1034. PMC 7889174. PMID 33435791.
- ^ Burki, Fabien; Roger, Andrew J.; Brown, Matthew W.; Simpson, Alastair G. B. (2020-01-01). "The New Tree of Eukaryotes". Trends in Ecology & Evolution. 35 (1): 43–55. doi:10.1016/j.tree.2019.08.008. ISSN 0169-5347. PMID 31606140. S2CID 204545629.
- ^ Linzhou Li; Sibo Wang; Hongli Wang; Sunil Kumar Sahu; Birger Marin; Haoyuan Li; Yan Xu; Hongping Liang; Zhen Li; Shifeng Chen; Tanja Reder; Zehra Çebi; Sebastian Wittek; Morten Petersen; Barbara Melkonian; Hongli Du; Huanming Yang; Jian Wang; Gane Ka-Shu Wong; Xun Xu; Xin Liu; Yves Van de Peer; Michael Melkonian; Huan Liu (22 June 2020). "The genome of Prasinoderma coloniale unveils the existence of a third phylum within green plants". Nature Ecology & Evolution. 4 (9): 1220–1231. doi:10.1038/s41559-020-1221-7. PMC 7455551. PMID 32572216.
- ^ Mackiewicz, Paweł; Gagat, Przemysław (2014-12-31). "Monophyly of Archaeplastida supergroup and relationships among its lineages in the light of phylogenetic and phylogenomic studies. Are we close to a consensus?". Acta Societatis Botanicorum Poloniae. 83 (4): 263–280. doi:10.5586/asbp.2014.044. ISSN 2083-9480.
- ^ de Vries, Jan; Archibald, John M. (2017). "Endosymbiosis: Did Plastids Evolve from a Freshwater Cyanobacterium?". Current Biology. 27 (3): R103–R105. doi:10.1016/j.cub.2016.12.006. PMID 28171752.
- ^ Lewis, L. A. (2017). "Hold the salt: Freshwater origin of primary plastids". Proceedings of the National Academy of Sciences of the United States of America. 114 (37): 9759–9760. doi:10.1073/pnas.1712956114. PMC 5604047. PMID 28860199.
- ^ Raven, John A. (2013). "Cells inside Cells: Symbiosis and Continuing Phagotrophy". Current Biology. 23 (12): R530–R531. doi:10.1016/j.cub.2013.05.006. PMID 23787050.
- ^ Andersson, Jan O.; Roger, Andrew J. (2002). "A cyanobacterial gene in non-photosynthetic protists – an early chloroplast acquisition in eukaryotes?". Current Biology. 12 (2): 115–119. doi:10.1016/S0960-9822(01)00649-2. PMID 11818061. S2CID 18809784.
- ^ Keeling, Patrick J. (2010). "The endosymbiotic origin, diversification and fate of plastids". Philosophical Transactions of the Royal Society B: Biological Sciences. 365 (1541): 729–748. doi:10.1098/rstb.2009.0103. PMC 2817223. PMID 20124341.
- ^ Bengtson, Stefan; Sallstedt, Therese; Belivanova, Veneta; Whitehouse, Martin (2017). "Three-dimensional preservation of cellular and subcellular structures suggests 1.6 billion-year-old crown-group red algae". PLOS Biology. 15 (3): e2000735. doi:10.1371/journal.pbio.2000735. PMC 5349422. PMID 28291791.
- ^ Javaux, Emmanuelle J.; Knoll, Andrew H.; Walter, Malcolm R. (2004). "TEM evidence for eukaryotic diversity in mid-Proterozoic oceans". Geobiology. 2 (3): 121–132. doi:10.1111/j.1472-4677.2004.00027.x. S2CID 53600639.
- ^ Yoon, Hwan Su; Hackett, Jeremiah D.; Ciniglia, Claudia; Pinto, Gabriele; Bhattacharya, Debashish (2004). "A molecular timeline for the origin of photosynthetic eukaryotes". Molecular Biology and Evolution. 21 (5): 809–818. doi:10.1093/molbev/msh075. PMID 14963099.
- ^ Butterfield, Nicholas J. (2000). "Bangiomorpha pubescens n. gen., n. sp.: implications for the evolution of sex, multicellularity, and the Mesoproterozoic/Neoproterozoic radiation of eukaryotes". Paleobiology. 26 (3): 386–404. doi:10.1666/0094-8373(2000)026<0386:BPNGNS>2.0.CO;2. S2CID 36648568.