Jakobea: diferenças entre revisões
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{{Automatic taxobox |
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#redirect[[Loukozoa]] |
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| image = Jakobida.svg |
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| image_caption = Four jakobid species, showing groove and flagella: ''[[Jakoba|Jakoba libera]]'' (ventral view), ''[[Stygiella (protist)|Stygiella incarcerata]]'' (ventral view), ''[[Reclinomonas|Reclinomonas americana]]'' (dorsal view), and ''[[Histiona|Histiona aroides]]'' (ventral view) |
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| taxon = Jakobida |
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| authority = Cavalier-Smith 1993 |
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| parent_authority = Cavalier-Smith 1997 emend. 2003<ref name="Cavalier-Smith 2003">{{cite journal|vauthors=Cavalier-Smith T|title=The excavate protozoan phyla Metamonada Grassé emend. (Anaeromonadea, Parabasalia, Carpediemonas, Eopharyngia) and Loukozoa emend. (Jakobea, Malawimonas): their evolutionary affinities and new higher taxa|journal=International Journal of Systematic and Evolutionary Microbiology|volume=53|date=2003|pages=1741–1758|doi=10.1099/ijs.0.02548-0}}</ref> |
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| subdivision_ranks = Families |
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| subdivision = |
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*[[Andaluciidae]] |
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*[[Stygiellidae]] |
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*[[Moramonadidae]] |
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*[[Jakobidae]] |
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*[[Histionidae]] |
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| synonyms = |
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*Jacobea {{au|Cavalier-Smith 1997}}<ref name="Cavalier-Smith 1997">{{cite journal|vauthors=Cavalier-Smith T|title=Amoeboflagellates and Mitochondrial Cristae in Eukaryote Evolution: Megasystematics of the New Protozoan Subkingdoms Eozoa and Neozoa|journal=Archiv für Protistenkunde|volume=147|date=1997|pages=237–258}}</ref> |
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| display_parents = 4 |
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}} |
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'''Jakobea''' é um filo de organismos de vida livre. |
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are an [[order (biology)|order]] of free-living, [[Heterotroph|heterotrophic]], [[Flagellate|flagellar]] [[eukaryote]]s in the supergroup [[Excavata]]. They are small (less than 15 μm), and can be found in aerobic and anaerobic environments.<ref name=kelly1993>{{cite journal |last1=O'Kelly |first1=Charles J. |title=The Jakobid flagellates: structural features of Jakoba, Reclinomonas, and Histonia and implications for the early diversification of eukaryotes |journal=Journal of Eukaryotic Microbiology |date=1993 |volume=40 |issue=5 |pages=627–636|doi=10.1111/j.1550-7408.1993.tb06120.x |s2cid=85938682 }}</ref><ref name=strassert2016>{{cite journal |last1=Strassert |first1=Jürgen F. H. |last2=Tikhonenov |first2=Denis V. |last3=Pombert |first3=Jean-François |last4=Kolisko |first4=Martin |last5=Tai |first5=Vera |last6=Mylnikov |first6=Alexander P. |last7=Keeling |first7=Patrick J. |title=Moramonas marocensis gen. nov., sp. nov.: a jakobid flagellate isolated from desert soil with a bacteria-like, but bloated mitochondrial genome |journal=Open Biology |date=2016 |volume=6 |issue=2 |pages=150239 |pmc=4772810 |pmid=26887409 |doi=10.1098/rsob.150239 }}</ref><ref name=burger2013>{{cite journal |last1=Burger |first1=Gertraud |last2=Gray |first2=Michael W. |last3=Forget |first3=Lise |last4=Lang |first4=B. Franz |title=Strikingly Bacteria-Like and Gene-Rich Mitochondrial Genomes throughout Jakobid Protists |journal=Genome Biology and Evolution |date=2013 |volume=5 |issue=2 |pages=418–438 |doi=10.1093/gbe/evt008 |pmid=23335123 |pmc=3590771 }}</ref> The order Jakobida, believed to be [[Monophyly|monophyletic]], consists of only twenty species at present, and was classified as a group in 1993.<ref name=kelly1993 /><ref name=burger2013 /><ref name=simpsonbook2017>{{cite book |last1=Simpson |first1=Alastair G. B. |editor1-last=Archibald |editor1-first=John M. |editor2-last=Simpson |editor2-first=Alastair G. B. |editor3-last=Slamovits |editor3-first=Claudio H. |title=Handbook of the Protists |publisher=Springer, Cham |pages=973–1003 |chapter=Jakobids|doi=10.1007/978-3-319-28149-0_6 |year=2017 |isbn=978-3-319-28147-6 }}</ref> There is ongoing research into the [[Mitochondrial DNA|mitochondrial genomes]] of jakobids, which are unusually large and bacteria-like, evidence that jakobids may be important to the evolutionary history of [[eukaryotes]].<ref name=strassert2016 /><ref name=lara2006>{{cite journal |last1=Lara |first1=Enrique |last2=Chatzinotas |first2=Antonis |last3=Simpson |first3=Alastair G. B. |title=Andalucia (n. gen.)—the Deepest Branch Within Jakobids (Jakobida; Excavata), Based on Morphological and Molecular Study of a New Flagellate from Soil |journal=Journal of Eukaryotic Microbiology |date=2006 |volume=53 |issue=2 |pages=112–120 |doi=10.1111/j.1550-7408.2005.00081.x |pmid=16579813 |s2cid=19092265 }}</ref> |
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Molecular phylogenetic evidence suggests strongly that jakobids are most closely related to [[Heterolobosea]] (Percolozoa) and [[Euglenozoa]].<ref name="pmid19237557">{{cite journal |vauthors=Hampl V, Hug L, Leigh JW, Dacks JB, Lang BF, Simpson AG, Roger AJ |title=Phylogenomic analyses support the monophyly of Excavata and resolve relationships among eukaryotic "supergroups" |journal=[[Proceedings of the National Academy of Sciences|Proc. Natl. Acad. Sci. U.S.A.]] |volume=106 |issue=10 |pages=3859–64 |date=February 2009 |pmid=19237557 |doi=10.1073/pnas.0807880106 |pmc=2656170 |bibcode=2009PNAS..106.3859H|doi-access=free }}</ref> |
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==Structure and Biology== |
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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.<ref name=simpson2001>{{cite journal |last1=Simpson |first1=Alastair G. B. |last2=Patterson |first2=David J. |title=On Core Jakobids and Excavate Taxa: The Ultrastructure of Jakoba incarcerata |journal=Journal of Eukaryotic Microbiology |date=2001 |volume=48 |issue=4 |pages=480–492 |doi=10.1111/j.1550-7408.2001.tb00183.x |pmid=11456326 |s2cid=24042909 }}</ref> 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.<ref name=lara2006 /> |
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<ref name=simpson2001 /> The [[Cell nucleus|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 [[Crista|cristae]]. Food [[vacuoles]] are mostly located on the cell posterior, and in most jakobids the [[endoplasmic reticulum]] is distributed throughout the cell.<ref name=simpsonbook2017 /> |
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The [[Sessility (motility)|sessile]], [[Lorica (biology)|loricate]] [[Histionidae]] and occasionally free-swimming ''[[Jakoba libera]]'' ([[Jakobidae]]) have [[extrusomes]] under the dorsal membrane that are theorized to be defensive structures.<ref name=kelly1993 /><ref name=simpsonbook2017 /> |
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==Ecology== |
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Jakobids are widely dispersed, having been found in soil, freshwater, and marine habitats, but generally not common.<ref name=strassert2016 /><ref name=lara2006 /><ref name=simpsonbook2017 /><ref name=lara2007>{{cite journal |last1=Lara |first1=Enrique |last2=Berney |first2=Cedric |last3=Ekelund |first3=Flemming |last4=Harms |first4=Hauke |last5=Chatzinotas |first5=Antonis |title=Molecular comparison of cultivable protozoa from a pristine and a polycyclic aromatic hydrocarbon polluted site |journal=Soil Biology and Biochemistry |date=2007 |volume=39 |issue=1 |pages=139–148 |doi=10.1016/j.soilbio.2006.06.017 |url=http://doc.rero.ch/record/17573/files/Lara_Enrique_-_Molecular_comparison_of_cultivable_protozoa_20100322.pdf }}</ref> However, [[environmental DNA]] surveys suggest that Stygiellidae are abundant in anoxic marine habitats.<ref name=simpsonbook2017 /><ref name=panek2015>{{cite journal |last1=Pánek |first1=Tomáš |last2=Táborský |first2=Petr |last3=Pachiadaki |first3=Maria G. |last4=Hroudová |first4=Miluše |last5=Vlček |first5=Čestmir |last6=Edgcomb |first6=Virginia P. |last7=Čepička |first7=Ivan |title=Combined Culture-Based and Culture-Independent Approaches Provide Insights into Diversity of Jakobids, an Extremely Plesiomorphic Eukaryotic Lineage |journal=Frontiers in Microbiology |date=2015 |volume=6 |page=art. 1288 |doi=10.3389/fmicb.2015.01288 |pmid=26635756 |pmc=4649034 |doi-access=free }}</ref> 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.<ref name=simpsonbook2017 /> 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.<ref name=panek2015 /> |
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All known jakobids are heterotrophic suspension feeders.<ref name=strassert2016 /><ref name=simpsonbook2017 /> Their primary prey is generally considered to be bacteria, though one species has been observed eating extremely small (< 1 µm) eukaryotic cells.<ref name=burger2013 /><ref>{{cite journal |last1=Christaki |first1=Urania |last2=Vázquez-Domínguez |first2=Evaristo |last3=Courties |first3=Claude |last4=Lebaron |first4=Phillipe |title=Grazing impact of different heterotrophic nanoflagellates on eukaryotic (''Ostreococcus tauri'') and prokaryotic picoautotrophs (''Prochlorococcus'' and ''Synechococcus'') |journal=Environmental Microbiology |date=2005 |volume=7 |issue=8 |pages=1200–1210 |doi=10.1111/j.1462-2920.2005.00800.x |pmid=16011757 }}</ref> Jakobids are generally slow swimmers, with low clearance rates relative to similar organisms.<ref name=simpsonbook2017 /> |
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No study has suggested jakobids might be pathogenic or toxic.<ref name=simpsonbook2017 /> |
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==Mitochondrial DNA== |
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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.<ref name=simpsonbook2017 /> |
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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.<ref name=burger2013 /><ref name=simpsonbook2017 /> 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.<ref name=simpsonbook2017 /> 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.<ref name=rodriguez2007>{{cite journal |last1=Rodriguez-Ezpeleta |first1=Naiara |last2=Brinkmann |first2=Henner |last3=Burger |first3=Gertraud |last4=Roger |first4=Andrew J. |last5=Gray |first5=Michael W. |last6=Philippe |first6=Herve |last7=Lang |first7=B. Franz |title=Toward Resolving the Eukaryotic Tree: The Phylogenetic Positions of Jakobids and Cercozoans |journal=Current Biology |date=2007 |volume=17 |issue=16 |pages=1420–1425 |doi=10.1016/j.cub.2007.07.036 |pmid=17689961 |doi-access=free }}</ref> |
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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.<ref name=simpsonbook2017 /> |
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Another hypothesis is that the phage-type RNA polymerase moved from one eukaryote group to another via [[Horizontal gene transfer|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.<ref name=simpsonbook2017 /> |
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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.<ref name=simpsonbook2017 /> However, the gene arrangement of jakobid mitochondrial DNA suggests an ancestral origin of bacteria-type RNA polymerase over a more-recent divergence.<ref name=burger2013 /><ref name=simpsonbook2017 /> |
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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.<ref name=simpsonbook2017 /><ref>{{cite journal |last1=Stechmann |first1=Alexandra |last2=Cavalier-Smith |first2=Thomas |title=Rooting the Eukaryote Tree by Using a Derived Gene Fusion |journal=Science |date=2002 |volume=297 |issue=5578 |pages=89–91 |pmid=12098695 |doi=10.1126/science.1071196 |bibcode=2002Sci...297...89S |s2cid=21064445 }}</ref> 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.<ref name=simpsonbook2017 /> |
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==Taxonomia== |
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Jakobida contains five families consisting of mostly free-swimming genera: [[Jakobidae]], [[Moramonadidae]], [[Andaluciidae]], and [[Stygiellidae]].<ref name=simpsonbook2017 /> The sixth family, [[Histionidae]], is largely populated by [[Sessility (motility)|sessile]] [[Lorica (biology)|loricate]] genera, and includes the first jakobids ever described.<ref name=simpsonbook2017 /> |
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Jakobids are a [[Monophyly|monophyletic]] group, and are most closely related to the [[Euglenozoa]] and [[Heterolobosea]].<ref name=burger2013 /><ref name=simpsonbook2017 /><ref name=rodriguez2007 /> |
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{{cladogram |
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|title=Cladogram of Jakobida<ref>{{Cite journal |last1=Yabuki |first1=Akinori |last2=Gyaltshen |first2=Yangtsho |last3=Heiss |first3=Aaron A. |last4=Fujikura |first4=Katsunori |last5=Kim |first5=Eunsoo |title=Ophirina amphinema n. gen., n. sp., a New Deeply Branching Discobid with Phylogenetic Afnity to Jakobids |journal=Scientific Reports |volume=8 |issue=16219 |doi=10.1038/s41598-018-34504-6 |year=2018 |page=16219 |pmid=30385814 |pmc=6212452 |bibcode=2018NatSR...816219Y |doi-access=free }}</ref> |
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|{{clade| style=font-size:90%;line-height:80% |
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|1={{Clade |
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|label1=Ophirinina |
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|1={{Clade |
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|label1=[[Ophirinidae]] |
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|1=''[[Ophirinia]]'' |
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}} |
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|2={{Clade |
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|label1=Andaluciina |
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|1={{Clade |
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|label1=[[Andaluciidae]] |
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|1=''[[Andalucia]]'' |
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|label2=[[Stygiellidae]] |
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|2=''[[Stygiella]]'' |
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}} |
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|label2=Histionina |
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|2={{Clade |
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|label1=[[Moramonadidae]] |
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|1={{Clade |
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|1=''[[Moramonas]]'' |
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|2=''[[Seculamonas]]'' |
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}} |
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|2={{Clade |
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|label1=[[Jakobidae]] |
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|1=''[[Jakoba]]'' |
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|label2=[[Histionidae]] |
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|2={{Clade |
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|1=''[[Reclinomonas]]'' |
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|2=''[[Histiona]]'' |
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}} |
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}} |
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}} |
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}} |
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}} |
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}} |
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}} |
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* '''Class [[Jakobea]]''' <small>Cavalier-Smith 1999</small> |
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** '''Order [[Jakobida]]''' <small>Cavalier-Smith 1993</small> |
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*** '''Suborder [[Ophirinina]]''' <small>Yabuki et al. 2018</small> |
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**** '''Family [[Ophirinidae]]''' <small>Yabuki et al. 2018</small> |
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***** Genus ''[[Ophirina]]'' <small>Yabuki et al. 2018</small> |
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****** Species ''[[Ophirina amphinema]]'' <small>Yabuki et al. 2018</small> |
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*** '''Suborder [[Andaluciina]]''' <small>Cavalier-Smith 2013</small> |
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**** '''Family [[Andaluciidae]]''' <small>Cavalier-Smith 2013</small> |
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***** Genus ''[[Andalucia (genus)|Andalucia]]'' <small>Lara et al. 2006</small> |
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****** Species ''[[Andalucia godoyi]]'' <small>Lara et al. 2006</small> |
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**** '''Family [[Stygiellidae]]''' <small>Pánek, Táborský & Čepička 2015</small><ref name=panek2015 /> |
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***** Genus ''[[Velundella]]'' <small>Pánek, Táborský & Čepička 2015</small> |
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****** Species ''[[Velundella nauta|V. nauta]]'' <small>Pánek, Táborský & Čepička 2015</small> |
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****** Species ''[[Velundella trypanoides|V. trypanoides]]'' <small>Pánek, Táborský & Čepička 2015</small> |
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***** Genus ''[[Stygiella (protist)|Stygiella]]'' <small>Pánek, Táborský & Čepička 2015 non Bruand 1853</small> |
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****** Species ''[[Stygiella incarcerata|S. incarcerata]]'' <small>(Bernard, Simpson & Patterson 2000) Pánek, Táborský & Čepička 2015</small> [''Jakoba incarcerata'' <small>Bernard, Simpson & Patterson 2000</small>; ''[[Andalucia incarcerata]]'' <small>(Bernard, Simpson & Patterson 2000) Lara et al. 2006</small>] |
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****** Species ''[[Stygiella agilis|S. agilis]]'' <small>Pánek, Táborský & Čepička 2015</small> |
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****** Species ''[[Stygiella cryptica|S. cryptica]]'' <small>Pánek, Táborský & Čepička 2015</small> |
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****** Species ''[[Stygiella adhaerens|S. adhaerens]]'' <small>Pánek, Táborský & Čepička 2015</small> |
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*** '''Suborder [[Histonina]]''' <small>Cavalier-Smith 1993</small> |
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**** Species ?''[[Jakoba echidna]]'' <small>O'Kelly 1991</small> |
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**** '''Family [[Moramonadidae]]''' <small>Strassert et al. 2016</small> |
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***** Genus ''[[Moramonas]]'' <small>Strassert et al. 2016</small> |
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****** Species ''[[Moramonas marocensis]]'' <small>Strassert et al. 2016</small> |
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***** Genus ''[[Seculamonas]]'' <small>Marx et al. 2003 nomen nudum</small> |
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****** Species ''[[Seculamonas ecuadoriensis]]'' <small>Marx et al. 2003 nomen nudum</small> |
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**** '''Family [[Jakobidae]]''' <small>Patterson 1990</small> |
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***** Genus ''[[Jakoba]]'' <small>Patterson 1990</small> |
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****** Species ''[[Jakoba bahamiensis]]'' <small>Burger & Lang (indeitum)</small> |
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****** Species ''[[Jakoba libera]]'' <small>(Ruinen 1938) Patterson 1990</small> [''Cryptobia libera'' <small>Ruinen 1938</small>] |
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**** '''Family [[Histionidae]]''' <small>Flavin & Nerad 1993</small> |
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***** Genus ''[[Histiona]]'' <small>Voigt 1902</small> [''[[Zachariasia]]'' <small>Voigt 1901 non Lemmermann 1895</small>] |
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****** Species ?''[[Histiona planctonica|H. planctonica]]'' <small>Scourfield 1937</small> |
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****** Species ''[[Histiona aroides|H. aroides]]'' <small>Pascher 1943</small> |
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****** Species ''[[Histiona velifera|H. velifera]]'' <small>(Voigt 1901) Pascher 1943</small> [''Zachariasia velifera'' <small>Voigt 1901</small>; ''Histiona zachariasii'' <small>Voigt 1901 nom. illeg.</small>] |
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***** Genus ''[[Reclinomonas]]'' <small>Flavin & Nerad 1993</small> |
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****** Species ''[[Reclinomonas americana|R. americana]]'' <small>Flavin & Nerad 1993</small> |
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****** Species ''[[Reclinomonas campanula|R. campanula]]'' <small>(Penard 1921) Flavin & Nerad 1993</small> [''Histiona campanula'' <small>Penard 1921</small>; ''Stenocodon campanula'' <small>(Penard 1921) Pascher 1942</small>] |
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***** Genus ''[[Stenocodon]]'' <small>Pascher 1942</small> |
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****** Species ''[[Stenocodon epiplankton]]'' <small>Pascher 1942</small> |
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***** Genus ''[[Stomatochone]]'' <small>Pascher 1942</small> |
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****** Species ''[[Stomatochone infundibuliformis|S. infundibuliformis]]'' <small>Pascher 1942</small> |
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****** Species ''[[Stomatochone cochlear|S. cochlear]]'' <small>Pascher 1942</small> |
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****** Species ''[[Stomatochone excavata|S. excavata]]'' <small>Pascher 1942</small> |
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****** Species ''[[Stomatochone epiplankton|S. epiplankton]]'' <small>Pascher 1942</small> |
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--------------> |
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==Ver também== |
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*[[Loukozoa]] |
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==Referências== |
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{{Reflist}} |
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{{Taxonbar|from=Q1135350}} |
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{{Classificação Eukaryota}} |
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{{Vida na Terra}} |
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[[Categoria:Jakobea| ]] |
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[[Categoria:Protistas]] |
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Revisão das 15h44min de 26 de junho de 2023
| Jakobea | |
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| Four jakobid species, showing groove and flagella: Jakoba libera (ventral view), Stygiella incarcerata (ventral view), Reclinomonas americana (dorsal view), and Histiona aroides (ventral view) | |
Classificação científica 
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| Predefinição taxonomia em falta (fix): | Jakobida |
| Families | |
| Sinónimos | |
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Jakobea é um filo de organismos de vida livre.
Ver também
Referências
- ↑ 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
- ↑ 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