Holozoa

Holozoa (vanakreeka sõnadest ὅλος 'terve, kogu' ja ζῷον 'loom') on organismide klaad, millesse kuuluvad loomad ja nende lähimad üherakulised sugulased, kuid mitte seened ja kõik teised organismid.

Kokku on selles rühmas 1,5 miljonit puhtheterotroofsete organismide liiki, sealhulgas umbes 300 üherakulist liiki.

Sel on mitmesuguseid alamrühmi, nimelt loomad (Metazoa) ja protistid kaelusviburlased (Choanoflagellata), Filasterea, Pluriformea ja ihtüospoorid (Ichthyosporea).

Koos seente ja mõnede teiste rühmadega on Holozoa osa eukarüootide ülemrühmast tagaviburstest.

Choanofila on nimetus, mida varem kasutati rühma kohta, mille koosseis on sarnane nagu rühmal Holozoa, kuid seda ei soovitata kasutada, sest sellesse ei kuulu loomad ning see on seetõttu parafüleetiline.

Holozoa protistidel on otsustav roll üherakulistest eellastest hulkraksete organismide tekkimise evolutsiooniliste sammude mõistmisel. Hiljutised genoomiuuringud on heitnud valgust evolutsioonilistele suhetele Holozoa evolutsiooniliste liinide vahel, aidates taibata hulkraksuse päritolu.

Mõned loomade kivististe kandidaadid on ümber tõlgendatud Holozoa protistideks.

Tunnused

Koosseis

Holozoa on klaad, kuhu kuuluvad loomad ja nende lähimad sugulased ja nende ühine eellane, kuid ei kuulu seened. See on defineeritud fülogeneetilise nomenklatuuri põhimõtete järgi kui klaad, mis hõlmab kõik, kes on Homo sapiens 'i sugulased, kuid ei hõlma seent Neurospora crassa. Peale loomade kuuluvad sinna erineva morfoloogiaga protistide liinid, nagu kaelusviburlased, ihtüospoorid ja eraldi perekonnad Corallochytrium, Syssomonas, ja Tunicaraptor.

Kaelusviburlased umbes 250 liigiga on loomade lähimad sugulased. Need on vabalt elavad või kolooniaid moodustavad viburloomad, kes toituvad bakteritest, kasutades iseloomulikku mikrovillidest "kaelust". Kaelusviburlaste kaelus on väga sarnane käsnade kaelusviburrakkudega, mistõttu on 19. sajandist saadik oletatud nende sugulust käsnadega. Salapärane Proterospongia on näide koloonialise eluviisiga kaelusviburlasest, keda on peetud käsnade päritoluga seotud olevaks. Teiste Holozoa protistide sugulust hakati nägema alles 1990. aastatel.

Ihtüospoorid (Ichthyosporea, ka Mesomycetozoea) hõlmavad umbes nelikümmend liiki, peamiselt parasiidid ja kommensalistid. Nende partnerid on mitmesugused loomad inimestest ja kaladest kuni mereselgrootuteni.

Viited

↑ Adl SM, Bass D, Lane CE, Lukeš J, Schoch CL, Smirnov A, Agatha S, Berney C, Brown MW, Burki F, Cárdenas P, Čepička I, Chistyakova L, del Campo J, Dunthorn M, Edvardsen B, Eglit Y, Guillou L, Hampl V, Heiss AA, Hoppenrath M, James TY, Karnkowska A, Karpov S, Kim E, Kolisko M, Kudryavtsev A, Lahr DJG, Lara E, Le Gall L, Lynn DH, Mann DG, Massana R, Mitchell EAD, Morrow C, Park JS, Pawlowski JW, Powell MJ, Richter DJ, Rueckert S, Shadwick L, Shimano S, Spiegel FW, Torruella G, Youssef N, Zlatogursky V, Zhang Q (2019). "Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes". Journal of Eukaryotic Microbiology. 66 (1): 4–119. DOI:10.1111/jeu.12691. PMC 6492006. PMID 30257078.
↑ Hehenberger, Elisabeth; Tikhonenkov, Denis V.; Kolisko, Martin; Campo, Javier del; Esaulov, Anton S.; Mylnikov, Alexander P.; Keeling, Patrick J. (2017). "Novel Predators Reshape Holozoan Phylogeny and Reveal the Presence of a Two-Component Signaling System in the Ancestor of Animals". Current Biology. 27 (13): 2043–2050.e6. DOI:10.1016/j.cub.2017.06.006. PMID 28648822.
↑ Tikhonenkov DV, Mikhailov KV, Hehenberger E, Mylnikov AP, Aleoshin VV, Keeling PJ, et al. (2020). "New Lineage of Microbial Predators Adds Complexity to Reconstructing the Evolutionary Origin of Animals". Current Biology. 30 (22): 4500–4509. DOI:10.1016/j.cub.2020.08.061. PMID 32976804.
↑ Sebé-Pedrós A, Degnan B, Ruiz-Trillo I (2017). "The origin of Metazoa: a unicellular perspective". Nature Reviews Genetics. 18 (8): 498–512. DOI:10.1038/nrg.2017.21. PMID 28479598. S2CID 30709486.
↑ Steenkamp, Emma T.; Wright, Jane; Baldauf, Sandra L. (jaanuar 2006). "The Protistan Origins of Animals and Fungi". Molecular Biology and Evolution. 23 (1): 93–106. DOI:10.1093/molbev/msj011. PMID 16151185.
↑ Simpson AGB, Slamovits CH, Archibald JM (2017). "Chapter 1. Protist Diversity and Eukaryote Phylogeny". Archibald JM, Simpson AGB, Slamovits CH (toim-d). Handbook of the Protists (inglise). Kd 1 (2 ed.). Springer International Publishing. Lk 1–22. ISBN 978-3-319-28147-6.
↑ Brunet T, King N (2022). "The Single-Celled Ancestors of Animals: A History of Hypotheses". Herron MD, Conlin PL, Ratcliff WC (toim-d). The Evolution of Multicellularity. Evolutionary Cell Biology. CRC Press. Lk 251–278. DOI:10.1201/9780429351907-17. ISBN 9780429351907.
↑ Ragan, Mark A.; Goggin, C. Louise; Cawthorn, Richard J.; Cerenius, Lage; Jamieson, Angela V.C.; Plourde, Susan M.; Rand, Thomas G.; Söoderhäll, Kenneth; Gutell, Robin R. (15. oktoober 1996). "A novel clade of protistan parasites near the animal-fungal divergence". PNAS. 93 (21): 11907–11912. Bibcode:1996PNAS...9311907R. DOI:10.1073/pnas.93.21.11907. PMC 38157. PMID 8876236.
Ros-Rocher N, Pérez-Posada A, Michelle LM, Ruiz-Trillo I (veebruar 2021). "The origin of animals: an ancestral reconstruction of the unicellular-to-multicellular transition". Open Biol. 11 (2): 200359. DOI:10.1098/rsob.200359. hdl:10261/251922. PMID 33622103.
Shalchian-Tabrizi, Kamran; Minge, Marianne A.; Espelund, Mari; Orr, Russell; Ruden, Torgeir; Jakobsen, Kjetill S.; Cavalier-Smith, Thomas; Aramayo, Rodolfo (7. mai 2008). Aramayo, Rodolfo (toim). "Multigene phylogeny of choanozoa and the origin of animals". PLOS ONE. 3 (5): e2098. Bibcode:2008PLoSO...3.2098S. DOI:10.1371/journal.pone.0002098. PMC 2346548. PMID 18461162.
Cavalier-Smith T (mai 2013). "Early evolution of eukaryote feeding modes, cell structural diversity, and classification of the protozoan phyla Loukozoa, Sulcozoa, and Choanozoa". European Journal of Protistology. 49 (2): 115–178. DOI:10.1016/j.ejop.2012.06.001. PMID 23085100.
Strother, Paul K.; Brasier, Martin D.; Wacey, David; Timpe, Leslie; Saunders, Martin; Wellman, Charles H. (aprill 2021). "A possible billion-year-old holozoan with differentiated multicellularity". Current Biology. 31 (12): 2658–2665.e2. DOI:10.1016/j.cub.2021.03.051. PMID 33852871.
Parfrey, Laura Wegener; Lahr, Daniel J. G.; Knoll, Andrew H.; Katz, Laura A. (16. august 2011). "Estimating the timing of early eukaryotic diversification with multigene molecular clocks". Proceedings of the National Academy of Sciences of the United States of America. 108 (33): 13624–13629. Bibcode:2011PNAS..10813624P. DOI:10.1073/pnas.1110633108. PMC 3158185. PMID 21810989.
Torruella, Guifré; de Mendoza, Alex; Grau-Bové, Xavier; Antó, Meritxell; Chaplin, Mark A.; del Campo, Javier; Eme, Laura; Pérez-Cordón, Gregorio; Whipps, Christopher M. (21. september 2015). "Phylogenomics Reveals Convergent Evolution of Lifestyles in Close Relatives of Animals and Fungi". Current Biology (English). 25 (18): 2404–2410. DOI:10.1016/j.cub.2015.07.053. ISSN 0960-9822. PMID 26365255.{{cite journal}}: CS1 hooldus: tundmatu keel (link)
Lang BF, O'Kelly C, Nerad T, Gray MW, Burger G (2002). "The Closest Unicellular Relatives of Animals". Current Biology. 12 (20): 1773–1778. DOI:10.1016/S0960-9822(02)01187-9. PMID 12401173.
Suga H, Chen Z, de Mendoza A, Sebé-Pedrós A, Brown MW, Kramer E, Carr M, Kerner P, Vervoot M, Sánchez-Pons N, Torruella G, Derelle R, Manning G, Lang BF, Russ C, Haas BJ, Roger AJ, Nusbaum C, Ruiz-Trillo I (2013). "The Capsaspora genome reveals a complex unicellular prehistory of animals". Nature Communications. 4: 2325. Bibcode:2013NatCo...4.2325S. DOI:10.1038/ncomms3325. PMC 3753549. PMID 23942320.
Urrutia A, Mitsi K, Foster R, Ross S, Carr M, Ward GM, et al. (2022). "Txikispora philomaios n. sp., n. g., a micro-eukaryotic pathogen of amphipods, reveals parasitism and hidden diversity in Class Filasterea". Journal of Eukaryotic Microbiology. 69 (2): e12875. DOI:10.1111/jeu.12875. PMID 34726818. S2CID 240422937.
Huldtgren T, Cunningham JA, Yin C, Stampanoni M, Marone F, Donoghue PCJ, Bengtson S (2011). "Fossilized Nuclei and Germination Structures Identify Ediacaran "Animal Embryos" as Encysting Protists". Science. 334 (6063): 1696–1699. Bibcode:2011Sci...334Q1696H. DOI:10.1126/science.1209537. PMID 22194575. S2CID 39813961.

[Adl_2019-1] Adl SM, Bass D, Lane CE, Lukeš J, Schoch CL, Smirnov A, Agatha S, Berney C, Brown MW, Burki F, Cárdenas P, Čepička I, Chistyakova L, del Campo J, Dunthorn M, Edvardsen B, Eglit Y, Guillou L, Hampl V, Heiss AA, Hoppenrath M, James TY, Karnkowska A, Karpov S, Kim E, Kolisko M, Kudryavtsev A, Lahr DJG, Lara E, Le Gall L, Lynn DH, Mann DG, Massana R, Mitchell EAD, Morrow C, Park JS, Pawlowski JW, Powell MJ, Richter DJ, Rueckert S, Shadwick L, Shimano S, Spiegel FW, Torruella G, Youssef N, Zlatogursky V, Zhang Q (2019). "Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes". Journal of Eukaryotic Microbiology. 66 (1): 4–119. DOI:10.1111/jeu.12691. PMC 6492006. PMID 30257078.

[Hehenberger-2017-2] Hehenberger, Elisabeth; Tikhonenkov, Denis V.; Kolisko, Martin; Campo, Javier del; Esaulov, Anton S.; Mylnikov, Alexander P.; Keeling, Patrick J. (2017). "Novel Predators Reshape Holozoan Phylogeny and Reveal the Presence of a Two-Component Signaling System in the Ancestor of Animals". Current Biology. 27 (13): 2043–2050.e6. DOI:10.1016/j.cub.2017.06.006. PMID 28648822.

[ChoanoEvo-3] Tikhonenkov DV, Mikhailov KV, Hehenberger E, Mylnikov AP, Aleoshin VV, Keeling PJ, et al. (2020). "New Lineage of Microbial Predators Adds Complexity to Reconstructing the Evolutionary Origin of Animals". Current Biology. 30 (22): 4500–4509. DOI:10.1016/j.cub.2020.08.061. PMID 32976804.

[Sebé-Pedrós_et_al._2017-4] Sebé-Pedrós A, Degnan B, Ruiz-Trillo I (2017). "The origin of Metazoa: a unicellular perspective". Nature Reviews Genetics. 18 (8): 498–512. DOI:10.1038/nrg.2017.21. PMID 28479598. S2CID 30709486.

[Steenkamp-5] Steenkamp, Emma T.; Wright, Jane; Baldauf, Sandra L. (jaanuar 2006). "The Protistan Origins of Animals and Fungi". Molecular Biology and Evolution. 23 (1): 93–106. DOI:10.1093/molbev/msj011. PMID 16151185.

[Handbook_of_the_Protists_1-6] Simpson AGB, Slamovits CH, Archibald JM (2017). "Chapter 1. Protist Diversity and Eukaryote Phylogeny". Archibald JM, Simpson AGB, Slamovits CH (toim-d). Handbook of the Protists (inglise). Kd 1 (2 ed.). Springer International Publishing. Lk 1–22. ISBN 978-3-319-28147-6.

[Brunet_&_King_2022-7] Brunet T, King N (2022). "The Single-Celled Ancestors of Animals: A History of Hypotheses". Herron MD, Conlin PL, Ratcliff WC (toim-d). The Evolution of Multicellularity. Evolutionary Cell Biology. CRC Press. Lk 251–278. DOI:10.1201/9780429351907-17. ISBN 9780429351907.

[8] Ragan, Mark A.; Goggin, C. Louise; Cawthorn, Richard J.; Cerenius, Lage; Jamieson, Angela V.C.; Plourde, Susan M.; Rand, Thomas G.; Söoderhäll, Kenneth; Gutell, Robin R. (15. oktoober 1996). "A novel clade of protistan parasites near the animal-fungal divergence". PNAS. 93 (21): 11907–11912. Bibcode:1996PNAS...9311907R. DOI:10.1073/pnas.93.21.11907. PMC 38157. PMID 8876236.

[Ros-Rocher_2021-9] Ros-Rocher N, Pérez-Posada A, Michelle LM, Ruiz-Trillo I (veebruar 2021). "The origin of animals: an ancestral reconstruction of the unicellular-to-multicellular transition". Open Biol. 11 (2): 200359. DOI:10.1098/rsob.200359. hdl:10261/251922. PMID 33622103.

[pmid18461162-10] Shalchian-Tabrizi, Kamran; Minge, Marianne A.; Espelund, Mari; Orr, Russell; Ruden, Torgeir; Jakobsen, Kjetill S.; Cavalier-Smith, Thomas; Aramayo, Rodolfo (7. mai 2008). Aramayo, Rodolfo (toim). "Multigene phylogeny of choanozoa and the origin of animals". PLOS ONE. 3 (5): e2098. Bibcode:2008PLoSO...3.2098S. DOI:10.1371/journal.pone.0002098. PMC 2346548. PMID 18461162.

[Early_Evo_Euk-11] Cavalier-Smith T (mai 2013). "Early evolution of eukaryote feeding modes, cell structural diversity, and classification of the protozoan phyla Loukozoa, Sulcozoa, and Choanozoa". European Journal of Protistology. 49 (2): 115–178. DOI:10.1016/j.ejop.2012.06.001. PMID 23085100.

[Strother_2021-12] Strother, Paul K.; Brasier, Martin D.; Wacey, David; Timpe, Leslie; Saunders, Martin; Wellman, Charles H. (aprill 2021). "A possible billion-year-old holozoan with differentiated multicellularity". Current Biology. 31 (12): 2658–2665.e2. DOI:10.1016/j.cub.2021.03.051. PMID 33852871.

[Parfrey-2011-13] Parfrey, Laura Wegener; Lahr, Daniel J. G.; Knoll, Andrew H.; Katz, Laura A. (16. august 2011). "Estimating the timing of early eukaryotic diversification with multigene molecular clocks". Proceedings of the National Academy of Sciences of the United States of America. 108 (33): 13624–13629. Bibcode:2011PNAS..10813624P. DOI:10.1073/pnas.1110633108. PMC 3158185. PMID 21810989.

[Torruella-2015-14] Torruella, Guifré; de Mendoza, Alex; Grau-Bové, Xavier; Antó, Meritxell; Chaplin, Mark A.; del Campo, Javier; Eme, Laura; Pérez-Cordón, Gregorio; Whipps, Christopher M. (21. september 2015). "Phylogenomics Reveals Convergent Evolution of Lifestyles in Close Relatives of Animals and Fungi". Current Biology (English). 25 (18): 2404–2410. DOI:10.1016/j.cub.2015.07.053. ISSN 0960-9822. PMID 26365255.{{cite journal}}: CS1 hooldus: tundmatu keel (link)

[Lang_et_al._2002-15] Lang BF, O'Kelly C, Nerad T, Gray MW, Burger G (2002). "The Closest Unicellular Relatives of Animals". Current Biology. 12 (20): 1773–1778. DOI:10.1016/S0960-9822(02)01187-9. PMID 12401173.

[Suga_et_al._2013-16] Suga H, Chen Z, de Mendoza A, Sebé-Pedrós A, Brown MW, Kramer E, Carr M, Kerner P, Vervoot M, Sánchez-Pons N, Torruella G, Derelle R, Manning G, Lang BF, Russ C, Haas BJ, Roger AJ, Nusbaum C, Ruiz-Trillo I (2013). "The Capsaspora genome reveals a complex unicellular prehistory of animals". Nature Communications. 4: 2325. Bibcode:2013NatCo...4.2325S. DOI:10.1038/ncomms3325. PMC 3753549. PMID 23942320.

[Txikispora-17] Urrutia A, Mitsi K, Foster R, Ross S, Carr M, Ward GM, et al. (2022). "Txikispora philomaios n. sp., n. g., a micro-eukaryotic pathogen of amphipods, reveals parasitism and hidden diversity in Class Filasterea". Journal of Eukaryotic Microbiology. 69 (2): e12875. DOI:10.1111/jeu.12875. PMID 34726818. S2CID 240422937.

[Huldtgren_et_al._2011-18] Huldtgren T, Cunningham JA, Yin C, Stampanoni M, Marone F, Donoghue PCJ, Bengtson S (2011). "Fossilized Nuclei and Germination Structures Identify Ediacaran "Animal Embryos" as Encysting Protists". Science. 334 (6063): 1696–1699. Bibcode:2011Sci...334Q1696H. DOI:10.1126/science.1209537. PMID 22194575. S2CID 39813961.