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Эволюция Metazoa С Позиции Гипотезы Галлертоида Формирование (По Grasshoff, 1993) Системы Каналов

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Эволюция Metazoa С Позиции Гипотезы Галлертоида Формирование (По Grasshoff, 1993) Системы Каналов Э в о л ю ц и я M e t a z o a https://www.scientificamerican.com/ Возраст жизни Возраст Земли - 4.5 - 4,6 млрд лет Древнейшие из известных пород - цирконы из Западной Австралии (Джек Хилз), их возраст не менее 4,4 млрд лет В Гренландии обнаружены оса- дочные породы возрастом 3,76 млрд лет (формация Исуа) с углеродом органического происхождения Жизнь на Земле появилась как минимум 3,76 млрд лет назад (4,2 млрд л.н. ?) ( из разных источников) Находки структурных остатков древнейших ископаемых микроорганизмов Бактерии. Квебек, Канада. Возраст Цианобактерии. Западная 4,3(?) - 3,7 млрд лет Австралия. Возраст 3,5 млрд лет) Dodd et al., 2017 Schopf et al. 2017 doi:10.1038/nature21377 10.1073/pnas.1718063115 Бактерии. Месторождение Варравуна, Западная Австралия. Возраст 3,5 млрд Соколов, Федонкин, 1996 лет); Last universal common ancestor (LUCA) Weiss et al, 2016 https://doi.org/10.1371/journal.pgen.1007518 https://astrobiology.nasa.gov/ Первые эукариоты Согласно палеонтологическим данным клетки с эукариотными параметрами появились 1,6 - 2 млрд. л.н. Knoll et al., 2006 Возможно, появление эукариот совпадает по времени с повышением концентрации O2. Ранние эукариоты Микрофоссилии из Roper Group (Северная Австралия) - 1,5 млрд. л.н. The scale bar in a is 35 m for a and c; 10 m for b; 100 m for d; 15 m for e; and 40 m for f. Javaux, Knoll, Walter, 2001 doi:10.1038/35083562 Obazoa Archaeplastida Holozoa Holomycota Stramenopiles Breviatea S A R Alveolata Apusomonadida Rhizaria Centroplasthelida Haptista Haptophyta Amoebozoa Cryptista Cryptophyta Metamonadida Fornicata Parabasalia “Excavata” Adl S. M. et al. Revisions to the classification, nomenclature, and diversity of eukaryotes //Journal Discoba of Eukaryotic Microbiology. – Euglenozoa 2019. – Т. 66. – №. 1. – С. 4-119. doi:10.1111/jeu.12691 Heterolobozea Макросистема эукариот 2019 Obazoa: Archaeplastida: Diaphoretickes Apusomonadida Glaucophyta S A R: Breviatea Rhodophyceae Opisthokonta Chloroplastida Stramenopiles Amorphea Holozoa Chlorophyta Bigyra (в.ч. Choanoflagellata Streptophyta (в.ч. Opalinata) Metazoa) (в.ч. Charophyceae Gyrista Amoebozoa: Nucletmycea Embryophyta) (в.ч. Oomycota Tubulinea в.ч. Fungi: Cryptista (?) Chrysophyceae Microsporidia (в.ч. Amoeba, Arcella, Phaeophyceae Chytridiomycota Difflugia) Xanthophyceae Dicaria Diatomista) Evosea Mucoromycota (в.ч. Eumycetozoa и др. Alveolata Archamoebida Colpodellida (в.ч. Pelomyxa, Perkinsidae Entamoeba)) Colponemida Discosea Dinoflagellata ? Apicomplexa Ciliophora Metamonada: ? Система Fornicata Rhiziaria (в.ч. Diplomonadida) Discoba: Cercozoa эукариот Parabasalia Jakobida Endomyxa (в.ч. Trichomonadida Tsukubamonadida Retaria (2019 г.) Spirotrichonymphida Heterolobosea (в.ч. Foraminifera Trichonymphida) Euglenozoa Radiolaria) Preaxostyla По Adl et al, 2019 (в.ч. Euglenida Aquavolonida doi:10.1111/jeu.12691 Группы неопределенного Kinetoplastea) положения Haptista (?) Переход к многоклеточности – одно из направлений эволюции эукариот Многократное возникновение многоклеточности у эукариот (филогенетическая схема упрощена) “In general, early-divergent persistent lineages are dominated by unicellular species (e.g., prasinophytes in the chlorobiontic clade), whereas later-divergent lineages contain a mixture of body plans (e.g., chlorophycean and charophycean algae). Species-rich, late-divergent persistent lineages tend to be exclusively multicellular (e.g., the land plants and metazoans)” doi:10.3732/ajb.1300314 Karl J. Niklas, 2014. American Journal of Botany 101(1): 6–25 Примеры многоклеточных протистов Volvox sp. Proterospongia sp. Dictyostelium sp. Колонии Псевдоплазмодий Многоклеточ- ные слоевища водорослей Corallina sp. Fucus sp. Первые многоклеточные эукариоты Около 1,5 млрд лет назад - начинается эпоха макроскопических многоклеточных организмов. Первые многоклеточные эукариоты Grypania – возможно, многоклеточная водоросль (древнейшие находки датированы 2,1 млрд л.) http://palaeos.com/proterozoic/paleoproterozoic/paleoproterozoic.html http://www.evrimagaci.org/makale/323 Первые многоклеточные эукариоты Bicellum brasieri – многоклеточный организм с дифференцированными клетками без клеточных стенок. Возможно, древнейший представитель Holozoa 5 мкм 5 мкм (1 млрд л.н.) 10 мкм Strother et al, 2021 https://doi.org/10.1016/j.cub.2021.03.051 Принципы организации Metazoa • Эукариотная клетка • Многоклеточное тело • Специализация, дифференцировка клеток • Ткани как качественно новый уровень специализации клеток • Особые механизмы интеграции Остатки Metazoa - следы их жизнедеятельности (норы и следовые дорожки на поверхности осадка) и структурные остатки - появились в палеонтологической летописи в позднем протерозое – около менее 1 млрд лет назад. Obazoa: Archaeplastida: Diaphoretickes Apusomonadida Glaucophyta S A R: Breviatea Rhodophyceae Opisthokonta Chloroplastida Stramenopiles Amorphea Holozoa Chlorophyta Bigyra (в.ч. Choanoflagellata Streptophyta (в.ч. Opalinata) Metazoa) (в.ч. Charophyceae Gyrista Amoebozoa: Nucletmycea Embryophyta) (в.ч. Oomycota Tubulinea в.ч. Fungi: Cryptista (?) Chrysophyceae Microsporidia (в.ч. Amoeba, Arcella, Phaeophyceae Chytridiomycota Difflugia) Xanthophyceae Dicaria Diatomista) Evosea Mucoromycota (в.ч. Eumycetozoa и др. Alveolata Archamoebida Colpodellida (в.ч. Pelomyxa, Perkinsidae Entamoeba)) Colponemida Discosea Dinoflagellata ? Apicomplexa Ciliophora Metamonada: ? Система Fornicata Rhiziaria (в.ч. Diplomonadida) Discoba: Cercozoa эукариот Parabasalia Jakobida Endomyxa (в.ч. Trichomonadida Tsukubamonadida Retaria (2019 г.) Spirotrichonymphida Heterolobosea (в.ч. Foraminifera Trichonymphida) Euglenozoa Radiolaria) Preaxostyla По Adl et al, 2019 (в.ч. Euglenida Aquavolonida doi:10.1111/jeu.12691 Группы неопределенного Kinetoplastea) положения Haptista (?) Воротничковые жгутиконосцы (Choanoflagellida) – ближайшие родственники Metazoa Sandeep Ravindran PNAS 2016;113:12889-12890 Жгутик Микровилле воротничка https://news.utexas.edu/2011/05/17/sodium_nervous_systems Клетка Monosiga brevicollis http://www.cell.com/trends/cell-biology/fulltext/S0962-8924(12)00167-5 Колония The King Lab kinglab.berkeley.edu Sphaeroeca sp. Howard Hughes Medical Institute and the Department of Molecular and Cell Biology at UC Berkeley Сравнение воротничковых клеток Choanoflagellida и губок Строение Choanoflagellida (A, Salpingoeca rosetta) и хоаноцитов губок (B, Sycon coactum). Отмечены жгутики (fL), микровилле воротничка (mv), ядро (nu), пищеварительные вакуоли (fv). Комплекс мастигонем присутствует и у воротничковых жгутиконосцев (C, Salpingoeca amphoridium) и у хоаноцитов (D, Spongilla lacustris). (E) Сравнение ультраструктуры клеток воротничкового жгутиконосца и хоаноцита. В некоторых исследованиях отмечено наличие филоподий и у хоаноцитов. Распроложение микротрубочек корешкового аппарата у Coanoflagellida (F) и хаоноцитов (G) Thibaut Brunet and Nicole King The Origin of Animal Multicellularity and Cell Differentiation. Developmental Cell. V. 43, Is. 2, 2017, Pages 124-140 https://doi.org/10.1016/j.devcel.2017.09.016 Spongia Choanoflagellida Сравнение воротничковых клеток губок (А) и Choanoflagellida (В) Строение кинетид хоаноцитов губок (А) и Choanoflagellida (В): ag- axial granule; bf – basal foot; bmt – band of microtubules; c – centriole; cfl - central filament; dz – electron-dense zone; fb – fibrillar bridge; fl – flagellum; fr – fibrillar root; Ga - Golgi apparatus; Similar characters: k - kinetosome; • developed transition fibers lfr – lateral fibrillar root; • centriole orthogonal and connected to kinetosome by a fibrillar bridge lmt - lateral microtubules; Different character: n – nucleus; A) choanoflagellates have, choanocytes have not: rM- ring of MTOCs; • central filament in transition zone (unique structure for eukaryotes) • transverse plate is always present tf - transition fiber • radial microtubular roots organized in bands • electron dense ring or foci as MTOCs • centriole produces fibrillar root to Golgi apparatus B) choanocytes have, choanoflagellates have not: Igor R. Pozdnyakov et al., 2017 • axial granule instead of transverse plate Kinetid structure of choanoflagellates and choanocytes • kinetosome with fibrillar root to the nucleus of sponges does not support their close relationship. • typical metazoan MTOCs (foot and satellites) Protistology, 11 (4) 2017, Pages 124-140 • dark region in flagellar transition zone doi:10.21685/1680-0826-2017-11-4-6 Monosiga sp. Жгутик Выросты клетки Внутриклеточная локализация кадгеринов Monosiga brevicollis (Choanoflagellida), Строение клетки выявленная методом Monosiga sp. иммунофлюоресценции Кадгерины - трансмембранные белки, характерные компоненты механических клеточных контактов Metazoa Abedin, King, 2008 (G and H) Choanoeca flexa colonies rapidly and reversibly invert their curvature while maintaining contacts among neighboring cells. (G) Flagella-in colony inverts to the flagella-out orientation (H) Flagella-out colony reverts to the flagella-in orientation (I) Summary of the inversion and relaxation processes. Brunet et al, 2019 DOI: 10.1126/science.aay2346 Данные палеоихнологии: трудности интерпретации Предполагаемые следы рытья или движения многоклеточных организмов Средний протерозой, Поздний эдиакарий. Stáhpogieddi формация Mackenzie Mountains, Formation. Scale bars - 1 cm Канада Narbonne, Aitken, 1990 Mcilroy, Brassier 2016 Первые многоклеточные эукариоты: трудности интерпретации Horodyskia williamsii, не позднее 1020 млн л. н. (другие виды - 1.5 млрд. лет н.) , Зап. Австралия. Конические зооиды колонии Maetazoa прикреплены к общему столону? Следы
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