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Jakobid

Jakobids are an order of free-living, heterotrophic, flagellar eukaryotes in the supergroup Excavata. They are small (less than 15 μm), and can be found in aerobic and anaerobic environments.[3][4][5] The order Jakobida, believed to be monophyletic, consists of only twenty species at present, and was classified as a group in 1993.[3][5][6] There is ongoing research into the mitochondrial genomes of jakobids, which are unusually large and bacteria-like, evidence that jakobids may be important to the evolutionary history of eukaryotes.[4][7]

Jakobid
Four jakobid species, showing groove and flagella: Jakoba libera (ventral view), Stygiella incarcerata (ventral view), Reclinomonas americana (dorsal view), and Histiona aroides (ventral view)
Scientific classification
Domain: Eukaryota
Clade: Diphoda
Clade: Discoba
Class: Jakobea
Cavalier-Smith 1997[1] em. 2003[2]
Order: Jakobida
Cavalier-Smith 1993
Families
Synonyms
  • Jacobidea Cavalier-Smith 1993

Molecular phylogenetic evidence suggests strongly that jakobids are most closely related to Heterolobosea (Percolozoa) and Euglenozoa.[8]

Structure and Biology edit

Jakobids have two flagella, inserted in the anterior end of the cell, and, like other members of order Excavata, have a ventral feeding groove and associated cytoskeleton support.[9] The posterior flagella has a dorsal vane and is aligned within the ventral groove, where it generates a current that the cell uses for food intake.[7][9] The nucleus is generally in the anterior part of the cell and bears a nucleolus. Most known jakobids have one mitochondrion, again located anteriorly, and different genera have flattened, tubular, or absent cristae. Food vacuoles are mostly located on the cell posterior, and in most jakobids the endoplasmic reticulum is distributed throughout the cell.[6]

The sessile, loricate Histionidae and occasionally free-swimming Jakoba libera (Jakobidae) have extrusomes under the dorsal membrane that are theorized to be defensive structures.[3][6]

Ecology edit

Jakobids are widely dispersed, having been found in soil, freshwater, and marine habitats, but generally not common.[4][7][6][10] However, environmental DNA surveys suggest that Stygiellidae are abundant in anoxic marine habitats.[6][11] Some are capable of surviving hypersaline and anoxic environments, though the Histionids have only been found in freshwater ecosystems, where they attach themselves to algae or zooplankton.[6] Outside of obligate sessile species, many species of jakobids can attach temporarily to surfaces, using either of the two flagella or the cell body itself.[11]

All known jakobids are heterotrophic suspension feeders.[4][6] Their primary prey is generally considered to be bacteria, though one species has been observed eating extremely small (< 1 µm) eukaryotic cells.[5][12] Jakobids are generally slow swimmers, with low clearance rates relative to similar organisms.[6]

No study has suggested jakobids might be pathogenic or toxic.[6]

Mitochondrial DNA edit

Since jakobids have no current commercial use, most research into jakobids has focused on their evolutionary significance. The mitochondrial DNA of jakobids is the most bacteria-like of all known eukaryotic mitochondrial DNA, suggesting that jakobid mitochondrial genomes might approximate the ancestral mitochondrial genome.[6]

Jakobid mitochondrial DNA is substantially different from most other eukaryotes, especially in terms of the number of genes (nearly 100 in some species) and bacteria-like elements within their genomes.[5][6] Nine of the genes have never been found in eukaryotic mitochondrial DNA. Uniquely, jakobid mitochondrial genomes code for bacteria-type RNA polymerase, as opposed to typical eukaryotic mitochondrial RNA polymerase, referred to as “phage-type”, which appears to be viral in origin.[6] This does not necessarily mean that jakobids are basal to the phylogeny of eukaryotes. While jakobid mitochondria have genetic features that seem to have developed from bacteria, and apparently lack phage-type RNA, it is possible that other eukaryotic clades lost their bacterial features independently.[13]

Several proposed possibilities might explain the bacterial features of jakobid mitochondrial DNA. One is that jakobids diverged very early from the rest of the eukaryotes. This hypothesis depends on whether or not jakobids are indeed basal to all living eukaryotes, but there is no evidence yet to support that suggestion.[6]

Another hypothesis is that the phage-type RNA polymerase moved from one eukaryote group to another via lateral gene transfer, replacing the bacteria-type enzyme, and simply did not reach the jakobids. This would not depend on jakobids being basal to eukaryotes as a whole, but has not been widely studied.[6]

A third possibility is the reverse of the others, suggesting that the phage-type RNA polymerase is the basal one. Under this scenario, jakobids acquired their bacteria-type RNA polymerase much more recently and that then spread via lateral gene transfer.[6] However, the gene arrangement of jakobid mitochondrial DNA suggests an ancestral origin of bacteria-type RNA polymerase over a more-recent divergence.[5][6]

One of the proposed scenarios suggests that the common ancestor of eukaryotes had two mitochondrial RNA polymerases, both phage-type and bacteria-type, and jakobids lost their phage-type polymerase while the rest of the eukaryotes lost the bacteria-type, possibly several times.[6][14] Such a model eliminates the need for jakobids to be truly basal. One study proposed that the phage-type and bacteria-type polymerases, when present in the same mitochondrion, served different functions, much in the way that the organelles of land plants have two different RNA polymerase enzymes that transcribe different genes.[6]

Taxonomy edit

Jakobida contains five families consisting of mostly free-swimming genera: Jakobidae, Moramonadidae, Andaluciidae, and Stygiellidae.[6] The sixth family, Histionidae, is largely populated by sessile loricate genera, and includes the first jakobids ever described.[6]

Jakobids are a monophyletic group, and are most closely related to the Euglenozoa and Heterolobosea.[5][6][13]

Cladogram of Jakobida[15]
Ophirinina
Ophirinidae

Ophirinia

Andalucina
Histionina
Moramonadidae

Moramonas

Seculamonas

  • Class Jakobea Cavalier-Smith 1999
    • Order Jakobida Cavalier-Smith 1993
      • Suborder Ophirinina Yabuki et al. 2018
        • Family Ophirinidae Yabuki et al. 2018
          • Genus Ophirina Yabuki et al. 2018
            • Species O. amphinema Yabuki et al. 2018
      • Suborder Andalucina Cavalier-Smith 2013
        • Family Andaluciidae Cavalier-Smith 2013
          • Genus Andalucia Lara et al. 2006
            • Species A. godoyi Lara et al. 2006
        • Family Stygiellidae Pánek, Táborský & Čepička 2015[11]
          • Genus Velundella Pánek, Táborský & Čepička 2015
            • Species V. nauta Pánek, Táborský & Čepička 2015
            • Species V. trypanoides Pánek, Táborský & Čepička 2015
          • Genus Stygiella Pánek, Táborský & Čepička 2015 non Bruand 1853
            • Species S. incarcerata (Bernard, Simpson & Patterson 2000) Pánek, Táborský & Čepička 2015 [Jakoba incarcerata Bernard, Simpson & Patterson 2000; Andalucia incarcerata (Bernard, Simpson & Patterson 2000) Lara et al. 2006]
            • Species S. agilis Pánek, Táborský & Čepička 2015
            • Species S. cryptica Pánek, Táborský & Čepička 2015
            • Species S. adhaerens Pánek, Táborský & Čepička 2015
      • Suborder Histonina Cavalier-Smith 1993
        • Species ?Jakoba echidna O'Kelly 1991
        • Family Moramonadidae Strassert et al. 2016
          • Genus Moramonas Strassert et al. 2016
            • Species M. marocensis Strassert et al. 2016
          • Genus Seculamonas Marx et al. 2003 nomen nudum
            • Species S. ecuadoriensis Marx et al. 2003 nomen nudum
        • Family Jakobidae Patterson 1990
          • Genus Jakoba Patterson 1990
            • Species J. bahamiensis Burger & Lang (indeitum)
            • Species J. libera (Ruinen 1938) Patterson 1990 [Cryptobia libera Ruinen 1938]
        • Family Histionidae Flavin & Nerad 1993
          • Genus Histiona Voigt 1902 [Zachariasia Voigt 1901 non Lemmermann 1895]
            • Species ?H. planctonica Scourfield 1937
            • Species H. aroides Pascher 1943
            • Species H. velifera (Voigt 1901) Pascher 1943 [Zachariasia velifera Voigt 1901; Histiona zachariasii Voigt 1901 nom. illeg.]
          • Genus Reclinomonas Flavin & Nerad 1993
            • Species R. americana Flavin & Nerad 1993
            • Species R. campanula (Penard 1921) Flavin & Nerad 1993 [Histiona campanula Penard 1921; Stenocodon campanula (Penard 1921) Pascher 1942]
          • Genus Stenocodon Pascher 1942
            • Species S epiplankton Pascher 1942
          • Genus Stomatochone Pascher 1942
            • Species S. infundibuliformis Pascher 1942
            • Species S. cochlear Pascher 1942
            • Species S. excavata Pascher 1942
            • Species S. epiplankton Pascher 1942

See also edit

References edit

  1. ^ Cavalier-Smith T (1997). "Amoeboflagellates and Mitochondrial Cristae in Eukaryote Evolution: Megasystematics of the New Protozoan Subkingdoms Eozoa and Neozoa". Archiv für Protistenkunde. 147: 237–258.
  2. ^ Cavalier-Smith T (2003). "The excavate protozoan phyla Metamonada Grassé emend. (Anaeromonadea, Parabasalia, Carpediemonas, Eopharyngia) and Loukozoa emend. (Jakobea, Malawimonas): their evolutionary affinities and new higher taxa". International Journal of Systematic and Evolutionary Microbiology. 53: 1741–1758. doi:10.1099/ijs.0.02548-0.
  3. ^ a b c O'Kelly, Charles J. (1993). "The Jakobid flagellates: structural features of Jakoba, Reclinomonas, and Histonia and implications for the early diversification of eukaryotes". Journal of Eukaryotic Microbiology. 40 (5): 627–636. doi:10.1111/j.1550-7408.1993.tb06120.x. S2CID 85938682.
  4. ^ a b c d Strassert, Jürgen F. H.; Tikhonenov, Denis V.; Pombert, Jean-François; Kolisko, Martin; Tai, Vera; Mylnikov, Alexander P.; Keeling, Patrick J. (2016). "Moramonas marocensis gen. nov., sp. nov.: a jakobid flagellate isolated from desert soil with a bacteria-like, but bloated mitochondrial genome". Open Biology. 6 (2): 150239. doi:10.1098/rsob.150239. PMC 4772810. PMID 26887409.
  5. ^ a b c d e f Burger, Gertraud; Gray, Michael W.; Forget, Lise; Lang, B. Franz (2013). "Strikingly Bacteria-Like and Gene-Rich Mitochondrial Genomes throughout Jakobid Protists". Genome Biology and Evolution. 5 (2): 418–438. doi:10.1093/gbe/evt008. PMC 3590771. PMID 23335123.
  6. ^ a b c d e f g h i j k l m n o p q r s t u Simpson, Alastair G. B. (2017). "Jakobids". In Archibald, John M.; Simpson, Alastair G. B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Springer, Cham. pp. 973–1003. doi:10.1007/978-3-319-28149-0_6. ISBN 978-3-319-28147-6.
  7. ^ a b c Lara, Enrique; Chatzinotas, Antonis; Simpson, Alastair G. B. (2006). "Andalucia (n. gen.)—the Deepest Branch Within Jakobids (Jakobida; Excavata), Based on Morphological and Molecular Study of a New Flagellate from Soil". Journal of Eukaryotic Microbiology. 53 (2): 112–120. doi:10.1111/j.1550-7408.2005.00081.x. PMID 16579813. S2CID 19092265.
  8. ^ Hampl V, Hug L, Leigh JW, Dacks JB, Lang BF, Simpson AG, Roger AJ (February 2009). "Phylogenomic analyses support the monophyly of Excavata and resolve relationships among eukaryotic "supergroups"". Proc. Natl. Acad. Sci. U.S.A. 106 (10): 3859–64. Bibcode:2009PNAS..106.3859H. doi:10.1073/pnas.0807880106. PMC 2656170. PMID 19237557.
  9. ^ a b Simpson, Alastair G. B.; Patterson, David J. (2001). "On Core Jakobids and Excavate Taxa: The Ultrastructure of Jakoba incarcerata". Journal of Eukaryotic Microbiology. 48 (4): 480–492. doi:10.1111/j.1550-7408.2001.tb00183.x. PMID 11456326. S2CID 24042909.
  10. ^ Lara, Enrique; Berney, Cedric; Ekelund, Flemming; Harms, Hauke; Chatzinotas, Antonis (2007). "Molecular comparison of cultivable protozoa from a pristine and a polycyclic aromatic hydrocarbon polluted site" (PDF). Soil Biology and Biochemistry. 39 (1): 139–148. doi:10.1016/j.soilbio.2006.06.017.
  11. ^ a b c Pánek, Tomáš; Táborský, Petr; Pachiadaki, Maria G.; Hroudová, Miluše; Vlček, Čestmir; Edgcomb, Virginia P.; Čepička, Ivan (2015). "Combined Culture-Based and Culture-Independent Approaches Provide Insights into Diversity of Jakobids, an Extremely Plesiomorphic Eukaryotic Lineage". Frontiers in Microbiology. 6: art. 1288. doi:10.3389/fmicb.2015.01288. PMC 4649034. PMID 26635756.
  12. ^ Christaki, Urania; Vázquez-Domínguez, Evaristo; Courties, Claude; Lebaron, Phillipe (2005). "Grazing impact of different heterotrophic nanoflagellates on eukaryotic (Ostreococcus tauri) and prokaryotic picoautotrophs (Prochlorococcus and Synechococcus)". Environmental Microbiology. 7 (8): 1200–1210. doi:10.1111/j.1462-2920.2005.00800.x. PMID 16011757.
  13. ^ a b Rodriguez-Ezpeleta, Naiara; Brinkmann, Henner; Burger, Gertraud; Roger, Andrew J.; Gray, Michael W.; Philippe, Herve; Lang, B. Franz (2007). "Toward Resolving the Eukaryotic Tree: The Phylogenetic Positions of Jakobids and Cercozoans". Current Biology. 17 (16): 1420–1425. doi:10.1016/j.cub.2007.07.036. PMID 17689961.
  14. ^ Stechmann, Alexandra; Cavalier-Smith, Thomas (2002). "Rooting the Eukaryote Tree by Using a Derived Gene Fusion". Science. 297 (5578): 89–91. Bibcode:2002Sci...297...89S. doi:10.1126/science.1071196. PMID 12098695. S2CID 21064445.
  15. ^ Yabuki, Akinori; Gyaltshen, Yangtsho; Heiss, Aaron A.; Fujikura, Katsunori; Kim, Eunsoo (2018). "Ophirina amphinema n. gen., n. sp., a New Deeply Branching Discobid with Phylogenetic Afnity to Jakobids". Scientific Reports. 8 (16219): 16219. Bibcode:2018NatSR...816219Y. doi:10.1038/s41598-018-34504-6. PMC 6212452. PMID 30385814.

jakobid, order, free, living, heterotrophic, flagellar, eukaryotes, supergroup, excavata, they, small, less, than, found, aerobic, anaerobic, environments, order, believed, monophyletic, consists, only, twenty, species, present, classified, group, 1993, there,. Jakobids are an order of free living heterotrophic flagellar eukaryotes in the supergroup Excavata They are small less than 15 mm and can be found in aerobic and anaerobic environments 3 4 5 The order Jakobida believed to be monophyletic consists of only twenty species at present and was classified as a group in 1993 3 5 6 There is ongoing research into the mitochondrial genomes of jakobids which are unusually large and bacteria like evidence that jakobids may be important to the evolutionary history of eukaryotes 4 7 JakobidFour jakobid species showing groove and flagella Jakoba libera ventral view Stygiella incarcerata ventral view Reclinomonas americana dorsal view and Histiona aroides ventral view Scientific classificationDomain EukaryotaClade DiphodaClade DiscobaClass JakobeaCavalier Smith 1997 1 em 2003 2 Order JakobidaCavalier Smith 1993FamiliesAndaluciidae Stygiellidae Moramonadidae Jakobidae HistionidaeSynonymsJacobidea Cavalier Smith 1993Molecular phylogenetic evidence suggests strongly that jakobids are most closely related to Heterolobosea Percolozoa and Euglenozoa 8 Contents 1 Structure and Biology 2 Ecology 3 Mitochondrial DNA 4 Taxonomy 5 See also 6 ReferencesStructure and Biology editJakobids have two flagella inserted in the anterior end of the cell and like other members of order Excavata have a ventral feeding groove and associated cytoskeleton support 9 The posterior flagella has a dorsal vane and is aligned within the ventral groove where it generates a current that the cell uses for food intake 7 9 The nucleus is generally in the anterior part of the cell and bears a nucleolus Most known jakobids have one mitochondrion again located anteriorly and different genera have flattened tubular or absent cristae Food vacuoles are mostly located on the cell posterior and in most jakobids the endoplasmic reticulum is distributed throughout the cell 6 The sessile loricate Histionidae and occasionally free swimming Jakoba libera Jakobidae have extrusomes under the dorsal membrane that are theorized to be defensive structures 3 6 Ecology editJakobids are widely dispersed having been found in soil freshwater and marine habitats but generally not common 4 7 6 10 However environmental DNA surveys suggest that Stygiellidae are abundant in anoxic marine habitats 6 11 Some are capable of surviving hypersaline and anoxic environments though the Histionids have only been found in freshwater ecosystems where they attach themselves to algae or zooplankton 6 Outside of obligate sessile species many species of jakobids can attach temporarily to surfaces using either of the two flagella or the cell body itself 11 All known jakobids are heterotrophic suspension feeders 4 6 Their primary prey is generally considered to be bacteria though one species has been observed eating extremely small lt 1 µm eukaryotic cells 5 12 Jakobids are generally slow swimmers with low clearance rates relative to similar organisms 6 No study has suggested jakobids might be pathogenic or toxic 6 Mitochondrial DNA editSince jakobids have no current commercial use most research into jakobids has focused on their evolutionary significance The mitochondrial DNA of jakobids is the most bacteria like of all known eukaryotic mitochondrial DNA suggesting that jakobid mitochondrial genomes might approximate the ancestral mitochondrial genome 6 Jakobid mitochondrial DNA is substantially different from most other eukaryotes especially in terms of the number of genes nearly 100 in some species and bacteria like elements within their genomes 5 6 Nine of the genes have never been found in eukaryotic mitochondrial DNA Uniquely jakobid mitochondrial genomes code for bacteria type RNA polymerase as opposed to typical eukaryotic mitochondrial RNA polymerase referred to as phage type which appears to be viral in origin 6 This does not necessarily mean that jakobids are basal to the phylogeny of eukaryotes While jakobid mitochondria have genetic features that seem to have developed from bacteria and apparently lack phage type RNA it is possible that other eukaryotic clades lost their bacterial features independently 13 Several proposed possibilities might explain the bacterial features of jakobid mitochondrial DNA One is that jakobids diverged very early from the rest of the eukaryotes This hypothesis depends on whether or not jakobids are indeed basal to all living eukaryotes but there is no evidence yet to support that suggestion 6 Another hypothesis is that the phage type RNA polymerase moved from one eukaryote group to another via lateral gene transfer replacing the bacteria type enzyme and simply did not reach the jakobids This would not depend on jakobids being basal to eukaryotes as a whole but has not been widely studied 6 A third possibility is the reverse of the others suggesting that the phage type RNA polymerase is the basal one Under this scenario jakobids acquired their bacteria type RNA polymerase much more recently and that then spread via lateral gene transfer 6 However the gene arrangement of jakobid mitochondrial DNA suggests an ancestral origin of bacteria type RNA polymerase over a more recent divergence 5 6 One of the proposed scenarios suggests that the common ancestor of eukaryotes had two mitochondrial RNA polymerases both phage type and bacteria type and jakobids lost their phage type polymerase while the rest of the eukaryotes lost the bacteria type possibly several times 6 14 Such a model eliminates the need for jakobids to be truly basal One study proposed that the phage type and bacteria type polymerases when present in the same mitochondrion served different functions much in the way that the organelles of land plants have two different RNA polymerase enzymes that transcribe different genes 6 Taxonomy editJakobida contains five families consisting of mostly free swimming genera Jakobidae Moramonadidae Andaluciidae and Stygiellidae 6 The sixth family Histionidae is largely populated by sessile loricate genera and includes the first jakobids ever described 6 Jakobids are a monophyletic group and are most closely related to the Euglenozoa and Heterolobosea 5 6 13 Cladogram of Jakobida 15 Ophirinina Ophirinidae OphiriniaAndalucina Andaluciidae AndaluciaStygiellidae StygiellaHistionina Moramonadidae MoramonasSeculamonasJakobidae JakobaHistionidae ReclinomonasHistionaClass Jakobea Cavalier Smith 1999 Order Jakobida Cavalier Smith 1993 Suborder Ophirinina Yabuki et al 2018 Family Ophirinidae Yabuki et al 2018 Genus Ophirina Yabuki et al 2018 Species O amphinema Yabuki et al 2018 Suborder Andalucina Cavalier Smith 2013 Family Andaluciidae Cavalier Smith 2013 Genus Andalucia Lara et al 2006 Species A godoyi Lara et al 2006 Family Stygiellidae Panek Taborsky amp Cepicka 2015 11 Genus Velundella Panek Taborsky amp Cepicka 2015 Species V nauta Panek Taborsky amp Cepicka 2015 Species V trypanoides Panek Taborsky amp Cepicka 2015 Genus Stygiella Panek Taborsky amp Cepicka 2015 non Bruand 1853 Species S incarcerata Bernard Simpson amp Patterson 2000 Panek Taborsky amp Cepicka 2015 Jakoba incarcerata Bernard Simpson amp Patterson 2000 Andalucia incarcerata Bernard Simpson amp Patterson 2000 Lara et al 2006 Species S agilis Panek Taborsky amp Cepicka 2015 Species S cryptica Panek Taborsky amp Cepicka 2015 Species S adhaerens Panek Taborsky amp Cepicka 2015 Suborder Histonina Cavalier Smith 1993 Species Jakoba echidna O Kelly 1991 Family Moramonadidae Strassert et al 2016 Genus Moramonas Strassert et al 2016 Species M marocensis Strassert et al 2016 Genus Seculamonas Marx et al 2003 nomen nudum Species S ecuadoriensis Marx et al 2003 nomen nudum Family Jakobidae Patterson 1990 Genus Jakoba Patterson 1990 Species J bahamiensis Burger amp Lang indeitum Species J libera Ruinen 1938 Patterson 1990 Cryptobia libera Ruinen 1938 Family Histionidae Flavin amp Nerad 1993 Genus Histiona Voigt 1902 Zachariasia Voigt 1901 non Lemmermann 1895 Species H planctonica Scourfield 1937 Species H aroides Pascher 1943 Species H velifera Voigt 1901 Pascher 1943 Zachariasia velifera Voigt 1901 Histiona zachariasii Voigt 1901 nom illeg Genus Reclinomonas Flavin amp Nerad 1993 Species R americana Flavin amp Nerad 1993 Species R campanula Penard 1921 Flavin amp Nerad 1993 Histiona campanula Penard 1921 Stenocodon campanula Penard 1921 Pascher 1942 Genus Stenocodon Pascher 1942 Species S epiplankton Pascher 1942 Genus Stomatochone Pascher 1942 Species S infundibuliformis Pascher 1942 Species S cochlear Pascher 1942 Species S excavata Pascher 1942 Species S epiplankton Pascher 1942See also editLoukozoaReferences edit Cavalier Smith T 1997 Amoeboflagellates and Mitochondrial Cristae in Eukaryote Evolution Megasystematics of the New Protozoan Subkingdoms Eozoa and Neozoa Archiv fur Protistenkunde 147 237 258 Cavalier Smith T 2003 The excavate protozoan phyla Metamonada Grasse emend Anaeromonadea Parabasalia Carpediemonas Eopharyngia and Loukozoa emend Jakobea Malawimonas their evolutionary affinities and new higher taxa International Journal of Systematic and Evolutionary Microbiology 53 1741 1758 doi 10 1099 ijs 0 02548 0 a b c O Kelly Charles J 1993 The Jakobid flagellates structural features of Jakoba Reclinomonas and Histonia and implications for the early diversification of eukaryotes Journal of Eukaryotic Microbiology 40 5 627 636 doi 10 1111 j 1550 7408 1993 tb06120 x S2CID 85938682 a b c d Strassert Jurgen F H Tikhonenov Denis V Pombert Jean Francois Kolisko Martin Tai Vera Mylnikov Alexander P Keeling Patrick J 2016 Moramonas marocensis gen nov sp nov a jakobid flagellate isolated from desert soil with a bacteria like but bloated mitochondrial genome Open Biology 6 2 150239 doi 10 1098 rsob 150239 PMC 4772810 PMID 26887409 a b c d e f Burger Gertraud Gray Michael W Forget Lise Lang B Franz 2013 Strikingly Bacteria Like and Gene Rich Mitochondrial Genomes throughout Jakobid Protists Genome Biology and Evolution 5 2 418 438 doi 10 1093 gbe evt008 PMC 3590771 PMID 23335123 a b c d e f g h i j k l m n o p q r s t u Simpson Alastair G B 2017 Jakobids In Archibald John M Simpson Alastair G B Slamovits Claudio H eds Handbook of the Protists Springer Cham pp 973 1003 doi 10 1007 978 3 319 28149 0 6 ISBN 978 3 319 28147 6 a b c Lara Enrique Chatzinotas Antonis Simpson Alastair G B 2006 Andalucia n gen the Deepest Branch Within Jakobids Jakobida Excavata Based on Morphological and Molecular Study of a New Flagellate from Soil Journal of Eukaryotic Microbiology 53 2 112 120 doi 10 1111 j 1550 7408 2005 00081 x PMID 16579813 S2CID 19092265 Hampl V Hug L Leigh JW Dacks JB Lang BF Simpson AG Roger AJ February 2009 Phylogenomic analyses support the monophyly of Excavata and resolve relationships among eukaryotic supergroups Proc Natl Acad Sci U S A 106 10 3859 64 Bibcode 2009PNAS 106 3859H doi 10 1073 pnas 0807880106 PMC 2656170 PMID 19237557 a b Simpson Alastair G B Patterson David J 2001 On Core Jakobids and Excavate Taxa The Ultrastructure of Jakoba incarcerata Journal of Eukaryotic Microbiology 48 4 480 492 doi 10 1111 j 1550 7408 2001 tb00183 x PMID 11456326 S2CID 24042909 Lara Enrique Berney Cedric Ekelund Flemming Harms Hauke Chatzinotas Antonis 2007 Molecular comparison of cultivable protozoa from a pristine and a polycyclic aromatic hydrocarbon polluted site PDF Soil Biology and Biochemistry 39 1 139 148 doi 10 1016 j soilbio 2006 06 017 a b c Panek Tomas Taborsky Petr Pachiadaki Maria G Hroudova Miluse Vlcek Cestmir Edgcomb Virginia P Cepicka Ivan 2015 Combined Culture Based and Culture Independent Approaches Provide Insights into Diversity of Jakobids an Extremely Plesiomorphic Eukaryotic Lineage Frontiers in Microbiology 6 art 1288 doi 10 3389 fmicb 2015 01288 PMC 4649034 PMID 26635756 Christaki Urania Vazquez Dominguez Evaristo Courties Claude Lebaron Phillipe 2005 Grazing impact of different heterotrophic nanoflagellates on eukaryotic Ostreococcus tauri and prokaryotic picoautotrophs Prochlorococcus and Synechococcus Environmental Microbiology 7 8 1200 1210 doi 10 1111 j 1462 2920 2005 00800 x PMID 16011757 a b Rodriguez Ezpeleta Naiara Brinkmann Henner Burger Gertraud Roger Andrew J Gray Michael W Philippe Herve Lang B Franz 2007 Toward Resolving the Eukaryotic Tree The Phylogenetic Positions of Jakobids and Cercozoans Current Biology 17 16 1420 1425 doi 10 1016 j cub 2007 07 036 PMID 17689961 Stechmann Alexandra Cavalier Smith Thomas 2002 Rooting the Eukaryote Tree by Using a Derived Gene Fusion Science 297 5578 89 91 Bibcode 2002Sci 297 89S doi 10 1126 science 1071196 PMID 12098695 S2CID 21064445 Yabuki Akinori Gyaltshen Yangtsho Heiss Aaron A Fujikura Katsunori Kim Eunsoo 2018 Ophirina amphinema n gen n sp a New Deeply Branching Discobid with Phylogenetic Afnity to Jakobids Scientific Reports 8 16219 16219 Bibcode 2018NatSR 816219Y doi 10 1038 s41598 018 34504 6 PMC 6212452 PMID 30385814 Retrieved from https en wikipedia org w index php title Jakobid amp oldid 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